Presented  in  honor  of 
Lawrence  C.  Spangard,  D.O* 


COLLEGE    OF    OSTEOPATHIC    PHYSICIANS 
AND  SURGEONS  •    LOS  ANGELES,  CALIFORNIA 

frlrlrlrlrlrMrlrlrlrlrT^^ 


BRAIN 
AND  SPINAL  CORD 

A  MANUAL -FOR  THE  STUDY  OF  THE  MORPHOLOGY  AND 
FIBRE-TRACTS  OF  THE  CENTRAL  NERVOUS  SYSTEM 


BY 

DR.  MED.  EMIL  VILL1GER 

PRIVATDOZENT  IN   NEUROLOGY  AND   NEUROPATHOLOGY  IN  THE   UNIVERSITY  OF  BASEL 


TRANSLATED  BY 

'  GEORGE  A.  PIERSOL,  M.D.,  Sc.D. 

PROFESSOR  OF  ANATOMY  IN  THE  UNIVERSITY  OF  PENNSYLVANIA 


SECOND  REVISED  AMERICAN  EDITION 

FROM  THE  THIRD  GERMAN  EDITION 

WITH  232  ILLUSTRATIONS 


PHILADELPHIA  $  LONDON 

J.  B.  LIPPINCOTT  COMPANY 


Copyright,  1912,  by  J.   B.  LIPPINCOTT  COMPANY 
Copyright,  1918,  by  J.  B.  LIPPINCOTT  COMPANY 


Electrotyped  and  Printed  by  J.  B.  Lippincott  Company, 
The  Washington  Square  Press,  Philadelphia,  U.  S.  A. 


\°\ 


TRANSLATOR'S  NOTE 


THE  increasing  attention  given  to  the  study  of  the  Central  Nervous  System  has 
emphasized  the  need  of  a  suitable  guide  for  laboratory  exercises.  The  usefulness  of 
Dr.  Villiger's  excellent  manual  has  been  greatly  increased  by  the  addition  of  Part  III, 
illustrating  the  architecture  of  the  brain-stem  by  series  of  consecutive  sections,  which 
first  appeared  in  the  second  edition.  While  of  much  assistance  tq  the  student  in  identi- 
fying details  under  the  microscope,  the  series  so  well  represents  the  actual  preparations, 
^  that  close  study  of  the  illustrations  alone  will  amply  repay  where  satisfactory  specimens 
are  inaccessible. 

The  translator  has  respected  the  author's  desire  to  retain  the  brevity  and  clearness 
which  characterize  the  book  ;  he  has  refrained,  therefore,  from  amplifying  the  text, 
•  which  appears,  with  slight  changes,  as  in  the  original.  Through  the  courtesy  of  the 
firm  of  Wilhelm  Engelmann,  of  Leipzig,  in  supplying  advance  proofs,  it  has  been  pos- 
sible to  include  the  new  figures,  which  have  been  added  to  the  third  German  edition. 
The  selected  bibliography,  appended  by  the  translator,  will  be  of  service,  it  is  hoped, 
to  those  desirous  of  consulting  the  original  papers  or  the  more  comprehensive  works 
pertaining  to  the  Central  Nervous  System. 

PHILADELPHIA, 

September,  1912. 


22802 


PREFACE  TO  THE  THIRD  EDITION 


THE  cordial  reception  accorded  the  second  edition  has  rendered  necessary,  within 
a  short  time,  issuing  a  new  edition.  Radical  changes  or  amplifications,  notably  the 
desired  additions  to  the  series  of  microscopical  representations  of  brain  sections,  could 
not  be  undertaken.  Notwithstanding  the  first  intention  to  print  the  edition  without 
alterations,  the  hearty  cooperation  of  the  publishers  has  made  it  possible  to  introduce, 
particularly  in  Part  II,  some  new  figures,  to  which  I  wish  to  direct  especial  attention. 

E.  VILLIGER. 
BASEL, 

April,  1912. 


AUTHOR'S    PREFACE 


THE  second  edition  presents  substantial  changes.  While  some  sections,  pertaining 
to  morphology  as  well  as  to  fibre-tracts,  have  been  simplified  and  made  clearer,  others 
have  been  treated  with  greater  completeness.  Certain  figures  have  been  replaced  by 
better  ones,  and  many  new  ones,  relating  to  the  paths  of  conduction,  have  been  added. 
The  chief  change,  however,  lies  in  the  addition  of  Part  III,  in  which  I  have  attempted 
to  meet  the  often  expressed  wish,  that  the  conduction-paths  be  represented  not  only  by 
diagrams,  but  also  by  microscopical  pictures.  I  am  well  aware  that  this  third  part 
presents  considerable  gaps.  Unfortunately  at  the  time  only  vertical  sections  through  the 
brain-stem  were  at  my  disposal ;  further,  the  drawing  of  the  new  and  especially  the 
microscopical  illustrations  made  such  claims  upon  me,  that  it  was  impossible  to  satisfy  all 
desires.  Nevertheless,  I  indulge  the  hope  that  the  study  of  the  fibre-tracts  will  be 
facilitated  by  the  microscopical  representations  now  given  and  by  the  newly  added 
schematic  figures. 

Moreover,  I  wish  particularly  to  emphasize,  that  this  second  edition  remains  a 
manual  and  as  such  is  designed  primarily  to  assist  the  student,  with  all  possible  con- 
ciseness and  clearness,  in  the  study  of  the  anatomy  of  the  central  nervous  system. 

It  is  my  privilege  to  take  this  opportunity  of  expressing  my  sincerest  thanks  to 
Professors  J.  Kollmann  and  N.  K.  Corning  for  their  kindness  in  placing  at  my  disposal 
numerous  microscopical  preparations.  My  especial  acknowledgment  is  due  the  firm  of 
Wilhelm  Engelmann,  of  Leipzig,  whose  cordial  cooperation  enabled  me  to  carry  out  the 
radical  changes  made  necessary  by  the  introduction  of  larger  figures,  particularly  the 

microscopical  illustrations. 

E.  VILLIGER. 


CONTENTS 


PART  I.— MORPHOLOGY. 


Subdivision  of  the  Central  Nervous  System. 

Development  of  the  Brain 

Development  of  the  Spinal  Cord 

Form,  Size  and  Weight  of  the  Brain 

General  Consideration  of  the  Brain 

Telencephalon— End- Brain 

Pallium — Cerebral  Mantle 

Rhinencephalon — Olfactory  Brain 

1.  Lobus  olfactorius 

A.  Lobus  olfactorius  anterior.  . . 

B.  Lobus  olfactorius  posterior . . . 

2.  Gyrus  fornicatus 

3.  Hippocampus 

4.  Gyrus  dentatus 

5.  Uncus  —  Gyrus  uncinatus.     Gyrus 

intralimbicus.       Gyrus     fascio- 
laris 

6.  Gyri  Andreae  Retzii 

Pars  optica  hypothalami 

Internal  Configuration 

Gray  Masses  and  Nuclei 

Summary  of  Telencephalon.  .• 

Diencephalon — Inter-Brain 

Thalamus  opticus 

Pars  mamillaris  hypothalami 

Ventriculus  tertius 

The  Nuclei  of  the  Diencephalon. . . . 
Summary 


PAGE 

Mesencephalon  —  Mid-Brain 

PAGE 

66 

3 
4 
.       8 

•       9 
.     ii 

Lamina  quadrigemina  
Pedunculi  cerebri  
Aquaeductus  cerebri  (Sylvii)  
Gray  Masses  of  the  Mid-Brain  
Summary  

.  .  .  .     66 
.  .  .  .     66 
.  .  .  .     68 
.  .  .  .     68 
69 

-     17 

Isthmus  rhombencephali 

7o 

.     18 

Metencephalon 

71 

•     25 
.     26 

Pons  Varolii  
Cerebellum 

.  ...     71 

72 

•     27 

A    Lobus  superior 

72 

-     29 

B    Lobus  posterior  

77 

•     30 

?2 

c.  Lobus  inferior  

.  .  .  .     74 

•     32 

s 

•     35 

Myelencephalon—  Medulla  oblongata  
Ventriculus  quartus  
Fossa  rhomboidea  
Gray  Masses  of  the  Rhombencephalon.  .  .  . 

.  .  .  .     77 
.  .  .  .     79 
.  ...     81 
.  .  .  .     82 
8s 

-      36 

Brain-Membranes  —  Meninges  . 

86 

•     37 

Dura  Mater            

-       87 

-     38 
.     46 

Arachnoidea.  ...     
Pia  Mater 

....     88 
88 

•     5° 

Spinal  Cord  —  Medulla  spinalis          .    .  . 

So 

•     52 
•     54 
•     56 

External  Configuration  
Internal  Configuration  
Membranes  of  the  Spinal  Cord     

....     89 
....     90 
q-j 

•     57 

Dura  mater  spinalis       

q-i 

•     58 

Arachnoidea  spinalis      *  

•  •  •  •      93 

.     61 

Pia  mater  sninalis.  .  . 

.      03 

PART  II.— FIBRE-TRACTS. 


Methods  of  Investigating  the  Fibre-Tracts  ....     97 

Histogenesis  of  the  Nervous  System 102 

Development  of  the  Ependyma  and  Neu- 

roglia  Cells 103 

Development  of  the  Nerve-Cells 105 

Development  of  the  Cells  of  the  Cerebro- 
Spinal  Ganglia  and  of  the  Sympathetic 

Ganglia 105 

The     Formed     Elements     of    the     Nervous 

System 106 

A.  The  Support-Cells     106 

B.  The  Nerve-Cells  107 

Microscopical    Structure    of     the     Cerebral 

Cortex 114 

I.  Cortex  of  the  Pallium 114 


II.  Rhinencephalon 

Bulbus  olfactorius 

Gyrus  fornicatus 

Hippocampus  and  Gyrus  dentatus. . 

Hippocampus 

Gyrus  dentatus 

Cerebral  Localization 

The  Motor  Centre 

The  Sensory  Centres — Sense-Centres . . 

The  Speech  Centres 

General  Division  of  the  Paths  of  Conduction . 
Conduction  Paths  of  the  Telencephalon 

1.  Association  Fibres 

2.  Commissural  Fibres 

3.  Projection  Fibres 


118 
118 
119 
1 20 

121 
122 

123 
124 
126 
128 
UI 
134 
134 
135 
135 


CONTENTS. 


A.  Short  Paths 135 

B.  Long  Paths 137 

Radiatio  corporis  striati 144 

Connections  of  the  Corpus  Striatum 144 

Fibre-Tracts  of  the  Rhinencephalon 144 

1.  Peripheral  Tract 144 

2.  Central  Tract 145 

A.  Connection  of    the    bulbus  olfac- 

torius       with      the       primary 
centres 145 

B.  Connection  of  the  primary  centres 

with   the    secondary  or  cortical 
centres '. 145 

3.  Connection  of  the  two  primary  centres  149 

4.  Further    connections     of    the    primary 

149 


of      the      two      cortical 


centres  . . . 

5.  Connection 

centres 149 

6.  Further    connections     of    the    cortical 

centres 149 

Conduction  Paths  of  the  Diencephalon 150 

Conduction  Paths  of  the  Mesencephalon 151 

Conduction  Paths  of  the  Metencephalon 154 

Microscopical  structure  of  the  Cerebellar 

Cortex  and  Fibre-Tracts 154 

Spinal  Cord 159 

The  Gray  Matter 159 

The  White  Matter...  ..  160 


1.  Paths  of  the  Anterior  Column   ....   161 

2.  Paths  of  the  Lateral  Column 161 

3.  Paths  of  the  Posterior  Column  ..    .   163 

Medulla  Oblongata 166 

Origin  of  the  Cerebral  Nerves 172 

Nervus   olfactorius 172 

Nervus  opticus 172 

Nervus  oculomotorius 175 

Nervus  trochlearis 176 

Nervus  abducens 176 

Nervus  trigeminus 176 

Nervus    facialis    and  Nervus  intermedius 

Wrisbergi 178 

Nervus  acusticus 179 

1.  Nervus  cochleae 179 

2.  Nervus  vestibuli 181 

Nervus  glossopharyngeus  and  vagus. ...     184 

Nervus  accessorius 185 

Nervus  hypoglossus 186 

General  Survey  of  the  Principal  Paths 186 

A.  Projection  Paths 186 

I.  Centripetal  Paths 186 

1.  Ascending  sensory  spinal  paths  186 

2.  Sensory  paths  of  the  cerebral 

nerves 1 88 

II.  Centrifugal  Paths 191 

B.  Reflex  Paths 192 

c.  Association  Paths 197 


PART  III.— SERIAL  SECTIONS  OF  THE  BRAIN-STEM. 


A.  From  the  anterior  end  of  the  corpus  cal- 

losum  to  the  quadrigeminal  region 205 


From  the  caudal  part  of  the  medulla  oblon- 
gata  to  the  quadrigeminal  region 235 


PART  I. 

MORPHOLOGY. 


BRAIN  AND  SPINAL  CORD 


MORPHOLOGY 


Medullary  plate 


Cnticle-plate 


Mesodertn 
Entoderm 


Chorda 
Medullary  groove 


--    Medullary  ridge 


The  brain  and  the  spinal  cord  together  constitute  the  central  nervous  system 
(sy ' sterna  nervorum  centrale^). 

The  brain  (encephalon)  is  that  part  of  the  central  nervous  system  lodged  within 
the  cranial  case ;  the  spinal  cord  (medulla  spinalis)  is  that  part  within  the  vertebral 
canal.  The  boundary  between  the  two  is  neither  macroscopically  nor  microscopically 
sharply  defined.  The  lowest  segment  of  the  brain  corresponds  perfectly  in  form  and 
structure  with  the  uppermost  one  of  the  spinal  cord 
and  is  called,  .therefore,  medulla  oblongata — the  length- 
ened marrow.  An  approximate  coarse  boundary  line 
is  supplied  by  the  lowest  bundle  of  the  so-called 
pyramidal  decussation,  or  by  the  highest  root-bundle 
of  the  first  cervical  nerve. 

A  further  separation  of  the  brain  into  different 
segments  is  best  accomplished  by  embryology.  The 
nervous  system  develops  from  a  broad  axial  stripe 
of  the  outer .  germ-layer,  the  ectoderm,  that  imme- 
diately overlies  the  chorda  dorsalis  or  noto  cord. 
Within  this  stripe,  the  cells  of  the  outer  germ-layer 
grow  into  elongated  cylindrical  or  spindle-form  elements, 
while  those  within  the  adjoining  ectoderm  become  flat- 
tened. In  this  manner  the  outer  germ-layer  differ- 
entiates into  tWO  zones:  (i)  the  thinned  OUt  Cuticle  FIG.  i.— Schematic  representation  of  the 

formation    of    the  medullary  tube  from    the 

plate    and    (2)  the    thicker   axially  placed    neural    or     outer  germ-layer. 
medullary  plate. 

The  two  zones  soon  become  more  sharply  defined  from  each  other;  the  medullary 
plate  curves  ventrally  and  at  its  margins  rises  above  the  surface  of  the  gern..  In  this 
manner  arise  the  medullary  ridges,  which  include  between  them  the  broad,  and  at  first 
shallow,  medullary  groove.  The  ridges  are  simple  folds  of  the  outer  germ-layer,  along 
the  juncture  of  the  medullary  and  cuticle  plates. 

3 


4  MORPHOLOGY. 

The  medullary  plates  are  converted  into  the  medullary  tube  very  early.  This 
tube  is  formed  by  a  typical  folding  process.  The  medullary  ridges  progressively  rise 
above  the  dorsal  surface  of  the  embryo,  bend  medially  and  grow  towards  each  other 
until  their  summits  meet  and  later  fuse.  As  the  medullary  ridges  rise  above  the  .surface 
of  the  embryonic  area,  they  draw  along  the  cuticle-plate ;  the  latter,  however,  does  not 
come  into  relation  with  the  nervous  system,  but  becomes  the  epithelial  covering  of  the 
body.  In  the  medullary  tube,  which  encloses  a  cleft-like  space,  the  central  canal  (can- 
alis  centralist,  filled  with  primary  lymph,  we  distinguish  the  brain-tube  and  the  spinal , 
tube ;  from  the  former  develops  the  brain  and  from  the  latter  the  spinal  cord. 


Anterior  brain-vesicle 
(  Prostncephalon) 


Middle  brain-vesicle 
(Mesencephalon) 


Pfsterior  brain-vesicle 
(Rhombencephalon) 


FIG.  2. — Schematic  representation  of  the  three  primary  brain-vesicles. 


DEVELOPMENT   OF   THE   BRAIN. 

The  fundamental  form  is  the  simple  brain-tube.  In  consequence  of  increased 
growth  in  certain  parts  and  diminished  growth  in  others,  the  brain-tube  early  exhibits  a 
segmentation.  At  first  it  consists  of  three  dilatations,  the  primary  brain-vesicles, 

separated  by  two  annu- 
lar constrictions,  the 
vesicles  being  desig- 
nated as  anterior,  mid- 
dle and  posterior.  From 
thes"e  three  brain-vesi- 
cles later  arise  the  three 
chief  divisions  :  the 
Fore-brain  or  Prosen- 
cephalon,  the  Mid-brain 
or  Mesencephalon  and  the  Hind-brain"  or  Rhombencephalon.  The  three  primary  vesicles 
subsequently  give  rise  to  five  secondary  brain-vesicles,  since  the  fore-brain  differen- 
tiates into  the  telencephalon  and  the  diencephalon,  while  the  hind-brain  divides  into 
the  metencephalon  and  the  myelen- 
cephalon.  The  hind-brain  is»  sepa- 
rated from  the  mid-brain  by  a  narrow 
constricted  segment,  the  isthmus 
(isthmus  rhombencephali) .  The  mye- 
lencephalon  is  continuous  with  the 
spinal  cord.  The  primitive  brain- 
tube,  therefore,  differentiates  into  six 
divisions  (Fig.  3)  :  the  telencephalon, 
the  diencephalon,  the  mesencephalon, 
the  isthmus,  the  metencephalon  and 
the  myelencephalon. 

In  the    later    stages,  the  devel- 
opment of   the  nervous   substance  is 

especially  vigorous  in  the  two  lateral  walls  of  the  neural  tube,  while  the  median  areas  of 
its  floor  and  roof  (the  floor-  and  roof-plates}  for  the  most  part  remain  thin  and  epithelioid. 
The  different  divisions  of  the  brain- tube  participate  in  the  further  development  in  very 


Telencephalo 


Myelencephalon 


FIG.  3. — The  five  secondary  brain-vesicles.     (His.) 


DEVELOPMENT  OF  BRAIN.  5 

unlike  degree.      Certain  segments  remain  far  behind,   while  others  far   outstrip  their  sur- 
roundings  in    consequence   of   their  vigorous    growth.      Along   with   the   displacement  of 


Telencephalon, -K& 


')iena/>ha.lon 


,.  Mesencepkalon 


Metencephalon 


Myelencephalo 


FIG,  4. — Brain  of  human  embryo  of  five  weeks.     After  a  model  by  His. 

certain  brain-segments  induced  by  unequal  growth,  other  processes  contribute  to  the  efface- 
ment   of  the  original  fundamental  plan  of  the  whole.     Among  such  factors  belong  partic- 


FIG.  5. — Brain  of  human  embryo  of  the  third  month.     After  a  model  by  His. 

ularly  the  appearance  of   robust  cross-fibres  (corpus  callosum,   pons) .      Consequently  it  is 
impossible  to  mark   off    superficially  the   individual  t  segments   on  the   brain  of   the  adult. 


6  MORPHOLOGY.  , 

The  developmental  relations  of  the  parts  of  the  brain  to  the  individual  brain-vesicles  is 
best  explained  by  the  accompanying  table  after  His.  It  will  serve  as  guide  in  the  con- 
sideration of  the  morphology.  (Compare  also  Figs.  5,  6,  7,  8,  9.) 


Pallium 


Telencephalon 


Cnrpus  strintnm 
Rhinencepkalon 


Pars  ofitica 
Hypvthalami 


FIG.  6. — Diagram  showing  the  further  development  of   the  fi 


The  prosencephalon  and    the   mesencephalon  together  are    also  designated  the  cere- 
brum or  great  brain.     The  brain-stem  (truncus  cerebri)  embraces  the  so-called  the  brain- 


FIG.  7.— Median  sagittal  section  through  the  adult    brain.     Telencephalon   is  yellow;    diencephalan  red;    mesencephalor 
blue;  metencephaloa  green;  myelencephalon  violet. 

ganglia;  it    consists    of   the    stem    of    the    end-brain,   the    inter-brain,   the  mid-brain,   the 
isthmus,  the  pons  and  the  medulla  oblongata. 


SUBDIVISIONS   OF  THE   BRAIN. 


Prosencephalon — Fore-Brain 


f  Pallium 
Hemisphaerium         ^   Rhinencephalon 

[  Stem  of  End- Brain 
[    Pars  optica  Hypothalami 
Pars  mamillaris  Hypothalami 


Telencephalon 


Diencephalon  f  Thalamus 

Thalamencephalon  j  Metathalamus 

[  Epithalamus 
Pedunculi  cerebri 


|      Mesencephalon-Mid-Brain      J  ........    { 

1    Corpora  quadrigemina 

I  f  Isthmus   rhombencephali 

I  Rhombencephalon—  Hind-Brain  j       Metencephalon       J    ^^  e 


Myelencephalon         Medulla  oblongata 


Corpus  callosum    (Trnncns) 

Sept.  pellucidnm 

Corpus  callosum  (Genn) 


Medulla  oblongata. 


FIG.  9. — Median  sagittal  section  through  the  brair 


8 


MORPHOLOGY. 


Foramen 

inter- 

/  ventricnlare 
(Monroi) 


The    cavities    of     the    embryonal    brain-vesicles    likewise    change    their    form    under 
the    influence    of   the.  various    growth-processes.       The    central    canal   of   the   spinal  cord 

is  continued  into  the  hind-part  of 
the  myelencephalon.  The  cavity  of 
the  fore -part  of  the  myelencephalon 
and  that  of  the  entire  metencephalon 
become  the  fourth  ventricle.  The 
cavity  of  the  mid-brain  remains  as 
the  aquaeductus  cerebri  or  Sylvian 
aqueduct.  The  cavity  of  the  dien- 
cephalon  or  inter -brain  becomes  the 
third  ventricle,  which  communicates 
with  the  lateral  ventricles  —  the  cav- 
ities of  the  hemisphere  -  vesicles  —  by 
means  of  the  Y-like  foramen  of  Monro 
(foramen  interventriculare} .  All  these 
spaces  are  filled  with  a  fluid,  the  liquor  cerebro-spinalis. 


FIG.  io.-Diagram  shoeing  the  brain-vent 


DEVELOPMENT   OF   THE   SPINAL   CORD. 

The  part  of  the  neural  tube  that  becomes  the  spinal  cord  appears  of  oval  form  on 
transverse  section.  The  central  canal  forms  a  dorso-ventrally  directed  cleft,  which  is 
bounded  laterally  by  the  thickened  walls  of  the  medullary  tube,  but  dorsally  and  ven- 
trally  by  thinner  parts  of  the  same  ;  therefore,  a  separation  into  a  right  and  left  half  is 


FIG.  ii. — Cross-section  of  spinal  cord  of  a 
human  embryo  of  four  and  one-half  weeks. 
(Sit.) 


FIG.   12. — Cross-section  of   spinal  cord  of  a  hu- 
man embryo  of  three  months.   (His.) 


easily  recognizable.  The  thinner  dorsal  and  ventral  walls  appear  as  commissures  behind 
and  before,  the  dorsal  or  posterior  commissure  being  called  the  roof-plate  and  the  ventral 
or  anterior  commissure  the  floor-plate.  During  the  further  development,  these  plates 
grow  relatively  little,  while  both  lateral  halves  continue  to  thicken,  their  growth  being 
especially  marked  ventrally.  In  this  locality  on  each  side  appears  a  ventral  projection. 
Consequently,  the  floor-plate  is  pushed  farther  from  the  surface  and,  finally,  a  median 
longitudinal  cleft,  the  fissura  mediana  anterior,  is  formed  in  front.  A  similar  change 


THE   SPINAL   CORD.  9 

occurs  in  the  dorsal  region,  the  roof-plate  being  likewise  pushed  in  aod  disappearing 
at  the  bottom  of  the  sulcus  medianus  posterior.  The  spinal  cord  now  consists  of  two 
robust  lateral  halves,  separated  from  each  other  by  an  anterior  fissure  and  a  posterior 
sulcus.  During  this  further  development  also  the  central  canal  "has  changed  its  form, 
since  the  dorsal  part  of  the  original  dorso-ventrally  directed  cleft  becomes  closed  in 
consequence  of  the  apposition  of  the  lateral  walls. 

At  first  the  spinal  cord  extends  the  entire  length  of  the  vertebral  canal  with  a  fairly 
constant  volume.  The  lower  end  of  the  cord  becomes  rudimentary  and  defined  from 
the  preceding  part,  assumes  a  conical  form  and  becomes  the  conus  medullaris.  A  fur- 
ther alteration  in  the  extension  of  the  spinal  cord  is  brought 
about  by  the  inequality  between  its  growth  and  that  of  the 
surrounding  vertebral  canal.  The  latter  constantly  increases 
in  length,  the  lower  segment  of  the  spine  developing  with 
especial  vigor.  Since  the  growth  of  the  cord  fails  to  keep 
pace  with  that  of  the  spine,  the  cord  apparently  shortens 
and  no  longer  extends  the  entire  length  of  the  vertebral 
canal.  The  conus  medullaris  is  drawn  up  from  the  sacral 
canal  and  enters  the  lumbar  region,  until,  finally,  it  is  found 
opposite  the  first  or  second  lumbar  vertebra.  During  this 
ascensus  medullae  spinalis  the  end  of  the  conus  medullaris 
is  drawn  out  into  a  thin  thread,  which  extends  as  far  as  the 
coccygeal  region  and  is  known  as  the  filum  terminate.  A 
further  consequence  of  this  ascensus  is  a  change  in  the 
course  of  the  nerves  emerging  from  the  spinal  cord.  In 
the  cervical  region  the  course  of  the  nerves  is  still  horizontal  ; 
in  the  thoracic  region  it  is  more  and  more  oblique  ;  while 
in  the  lumbar  region,  and,  still  more  in  the  sacral,  the  nerves 
are  directed  downward.  The  nerve-trunks  emerging  from 
the  last  part  of  the  cord  lie,  therefore,  for  a  long  distance 
within  the  vertebral  canal  before  they  leave  the  latter.  They 
surround  the  conus  medullaris  and  the  filum  terminale  and 
in  this  manner  lead  to  the  formation  of  the  so-called  horse-tail  or  cauda  equina.  In 
completion,  the  spinal  cord  undergoes  some  further  changes  in  its  form.  Gradually  two 
segments  acquire  greater  development,  the  one  in  the  cervical  portion  and  the  other  in 
the  upper  part  of  the  lumbar  region.  They  are  known  as  the  cervical  enlargement 
(intumescentia  cervicalis}  and  the  lumbar  enlargement  (intumescentia  lumbalis}  respectively. 


Conus 
medullaris 


Filum 
terminale 


FIG.   13. — Anterior   aspect    of   spinal 
cord.     Schematic. 


FORM,    SIZE   AND   WEIGHT   OF   THE   BRAIN. 

The  brain  possesses  in  a  general  way  the  form  of  the  cranial  cavity.  It  is 
applied  so  closely  to  the  inner  wall  of  the  skull,  that  a  cast  of  the  cranial  cavity 
repeats  to  a  considerable  degree  the  form  of  the  brain.  Corresponding  to  the  numerous 
variations  in  the  configuration  of  the  skull,  sometimes  the  brain  is  more  spherical 
and  at  other  times  more  ellipsoidal  in  form.  Its  dorsal  surface  is  arched,  its  ventral 
one  flattened. 


io  MORPHOLOGY. 

The  length  of  the  brain  is,  on  an  average,  between  160-170  mm.  and  its  greatest 
transverse  diameter  140  mm.  The  female  brain  is  usually  somewhat  shorter  than  the 
male.  The  weight  of  the  brain  has  long  been  the  subject  of  numerous  investigations. 
The  average  brain-weight  of  the  adult  man  has  been  found  to  be  1375  grams,  that  of 
the  adult  woman  1245  grams.  The  minimal  weight  of  the  male  brain  has  been  placed 
at  960  grams,  that  of  the  female  brain  at  800  grams.  As  maximal  weights,  2000 
grams  and  over,  1900  grams,  1861  grams  and  1807  grams  have  been  specified. 

The  difficulty  of  determining  the  average  brain-weight  lies  in  the  fact  that  various 
factors  exert  a  substantial  influence.  In  this  connection  age  plays  a  prominent  role. 
Observations  show  that  the  mean  brain-weight  in  both  sexes  reaches  its  maximum 
towards  the  twentieth  year,  remains  stationary  between  the  twentieth  and  fiftieth  years, 
and  then  gradually  decreases.  Further  influences  are  body-weight  and  body-length.  In 
general,  heavier  individuals  possess  a  heavier  brain  and  with  increase  in  height  is  asso- 
ciated increase  in  brain-weight ;  small  individuals,  however,  possess  a  relatively  heavier 
brain  than  large  ones.  In  relation  to  the  form  of  the  skull,  a  higher  average  brain- 
weight  has  been  found  in  the  broad-headed  type  than  in  the  long-headed.  Many  obser- 
vations regarding  the  influence  of  race  exist,  with  the  following  results  : 

Grams. 

Caucasian  race  :  average  brain-weight,  1 335 

.    Chinese:  average  brain-weight,  1332 

Sandwich  Islander:  average  brain-weight,  1303 

Malay  and  Indian  :  average  brain-weight,  1266 

Negro  :  average  brain-weight,  1244 

Australian:  average  brain- weight,  1185 

Definite  differences  in  the  brain-weight  among  the  European  nations  have  been 
recorded: 

Grams. 

German :  average  brain-weight,  1425 
English :  average  brain- weight,  1346 
French:  average  brain- weight,  1280 

Among  all  peoples,   the  female  sex  shows  a  smaller  average  brain-weight. 

Further,  the  influence  of  culture  is  to  be  noted.  According  to  the  measurements 
of  P.  Broca,  among  the  cultured  nations  the  brain-mass  probably  gains  somewhat  in  the 
course  of  time.  Based  on  the  measurements  of  Egyptian  skulls,  E.  Schmidt  found  that 
nations,  which  have  regressed  from  a  higher  culture,  exhibit  a  smaller  cranial  capacity 
than  that  possessed  by  them  during  the  period  of  their  cultural  bloom. 

Finally,  pathological  conditions  must  also  be  considered,  since  sometimes  they  in- 
duce an  increase  and  at  other  times  a  decrease  of  the  brain-weight. 

.  Of  great  interest  has  always  been  the  question,  to  what  extent  do  the  absolute  and 
relative  proportions  of  the  brain  indicate  the  favored  position  which  man  enjoys  in  com- 
parison with  other  animals.  It  has  long  been  known,  that  man  does  not  possess  abso- 
lutely the  heaviest  brain.  The  brain-weight  of  the  elephant  reaches  4000  grams  and 
more,  while  that  of  certain  cetaceans  may  be  3000  grams.  It  is,  however,  clear  that, 


BRAIN-WEIGHT.  n 

in  proportion  to  body-weight,  these  animals  possess  relatively  a  smaller  brain-mass  than 
does  man.  On  the  other  hand,  several  investigators  have  shown  that  man  does  not 
possess  relatively  the  heaviest  brain,  since  in  this  respect  he  is  surpassed  by  certain  song- 
birds, apes  and  mice.  .If,  however,  one  compares,  as  did  Ranke,  the  weight  of  the 
spinal  cord  with  that  of  the  brain,  man  is  found  to  possess  the  heaviest  brain.  While 
this  proportion  in  the  adult  human  subject  is  approximately  2  per  cent.,  in  the  anthro- 
poid apes  this  ratio  increases  to  about  6  per  cent.,  and  among  the  other  mammals  it 
rises  to  from  23  to  47  per  cent. 

It  is  particularly  difficult  to  establish  a  definite  relation  between  brain-weight  and 
intelligence.  The  comparison  of  many  brains  shows,  that  it  is  not  permissible  to  esti- 
mate the  intellectual  capacity  of  an  individual  merely  according  to  his  brain-weight.  The 
following  data  exist  regarding  the  weights  of  the  brains  of  distinguished  men: 


Turgenjeff :    2012  Broca:  1484 

Cuvier :  1861  Dupuytren :  1437 

Byron  :  1807  Dante  :  1420 

Kant:  1600  Liebig :  1352 

Schiller:        1580  Tiedemann :  1254 

Gauss:  J492  Dollinger:  1207 

This  comparison  shows,  that  while  the  majority  of  these  brains  exceeded  the 
average  weight  of  1375  grams,  there  are  also  men  of  eminent  intellect  who  possess  a 
relatively  low  brain-weight.  Moreover,  there  are  records  of  notable  brain-weights  (2028 
and  1900  grams)  among  individuals  of  insignificant  mentality.  Remarkably  lo*v  brain- 
weights  (300  grams  and  less)  occur  among  idiots. 

According  to  the  present  investigations,  the  conclusion  is  justified,  that  psychic 
functions  can  proceed  normally  only  where  the  brain-weight  has  passed  a  certain  mini- 
mum. According  to  Obersteiner,  the  lowest  level  to  which  the  brain-weight  may  sink 
without  noticeable  impairment  of  the  intellectual  faculties  is  for  the  male  brain  1000 
grams  and  for  the  female  900. 

It  is  to  be  noted,  that  weighing  the  entire  brain  supplies  only  an  uncertain  index 
of  the  psychic  capability,  for  the  reason  that  the  individual  parts  of  the  brain,  so  varying 
in  structure  and  function,  do  not  undergo  uniform  increase  or  diminution  in  size  and 
weight.  An  accurate  knowledge  of  the  weights  of  the  individual  parts  of  the  brain 
would  be  of  great  importance,  especially  an  exact  determination  of  the  weight  of  the 
gray  substance  of  the  end-brain,  the  cerebral  cortex,  with  which  particularly  the  higher 
psychic  functions  are  associated.  Even  then  we  would  fail  to  reach  a  positive  result, 
since,  in  addition  to  the  weight,  other  relations  must  be  considered,  especially  the  finer 
structure. 

GENERAL  INSPECTION  OF  THE  BRAIN. 

Let  us  first  examine  the  dorsal  surface  of  the  brain.  This  is  strongly  arched  in 
the  sagittal  as  well  as  in  the  frontal  direction— -fades  convexa  cerebri  (Fig.  14).  A 
deep  median  vertical  cleft  (fissura  longitudinalis  cerebri}  divides  the  whole  into  two 


12 


MORPHOLOGY. 


symmetrical  halves,   the  hemispheres  of  the  end-brain.      On  probing  to  the  bottom  of  the 
fissure,   one   learns  that   the   separation  is  not    complete,   since  in  the  middle  of  the   cleft 

the  two  halves  are  united  by  a  broad  hori- 
zontal commissure,  the  corpus  callosum.  In 
front  of  the  latter,  the  fissure  passes  to  the 
ventral  surface  of  the  brain ;  behind  the 
commissure,  the  fissure  likewise  penetrates 
deeply  and  ends  in  a  large  transverse  cleft 
(fissura  transversa  cerebri),  which  separates 
the  hemispheres  from  the  subjacent  cere- 
bellum. The  surface  of  the  hemispheres  ex- 
hibits clefts  and  furrows  of  varying  depths 
and  the  intervening  convolutions. 

The  ventral  surface  of  the  brain,  known 
as  the  basis  cerebri,  is  much  more  com- 
plexly modelled.  In  the  first  place,  we  per- 
ceive to  what  extent  the  hemispheres  occupy 
also  the  base  of  the  brain.  In  the  anterior 
part,  the  fissura  longitudinalis  cerebri  runs 
in  the  mid-line,  as  far  backward  as  an  X- 

PlG.  14.— Brain  viewed  from  above;  frontal  pole  below,     shaped     Structure,      the      chidSma      Oplicum.         On 


Bulbus  olfact. 


Fissiira  longitudinalis  ctrebri 


Tract,  olfact 


N.  access 


Cerebellum 


med.  ant. 

Incisura       Medulla       Sulc. 
cerebelli  post,      oblongata.      lateral,  ant. 


FIG.   15. — Basal  aspect  of  the  brain. 


BASAL  ASPECT  OF  BRAIN.  13 

folding  the  chiasma  slightly  backward,  one  sees  a  thin  gray  and  easily  torn  lamella 
stretching  from  the  front  border  of  the  chiasma  into  the  depth  of  the  fissura  longitudi- 
nalis  cerebri  ;  this  is  the  lamina  terminalis.  Forwards  from  the  chiasma  lead  the  nerves 
of  sight  (nervi  optici},  while  posteriorly  and  laterally,  on  each  side,  extends  the  visual 
path,  the  tractus  optici.  Lateral  from  the  chiasma  and  the  optic  tract  lies  a  gray  field, 
penetrated  by  larger  and  smaller  openings,  the  substantia  perforata  anterior.  The 
anterior  boundary  of  this  field  presents  a  triangular  area,  the  trigonum  olfactoriwn,  from 
whose  front  point  a  narrow  white-stripe,  the  tractus  olfactorius,  leads  forward  to  end  in 
the  broadened  terminal  bulbus  olfactorius.  The  olfactory  nerve-fibres  (fila  olfactoria) 
extend  from  the  ventral  surface  of  the  bulb  as  delicate  white  thread-like  strands,  that 
have  been  torn  in  removing  the  brain.  Bulbus  olfactorius,  tractus  olfactorius,  trigonum 
olfactorium,  substantia  perforata  anterior  are  all  parts  of  the  rhinencephalon.  These  will 
be  more  closely  considered  in  connection  with  the  rhinencephalon. 

Behind  the  chiasma  opticum  rises  a  gray  hump,  the  tuber  cinereum,  that  tapers  to 
the  infundibulum  bearing  a  bean-shaped  gray  body,  the  hypophysis  or  pituitary  body. 
The  hypophysis  lies  in  the  sella  turcica  of  the  body  of  the  sphenoid  and  may  readily 
become  separated  in  consequence  of  the  tearing  of  the  thin  infundibulum,  when  the 
brain  is  taken  out,  so  that  only  the  conical  pointed  part  of  the  infundibulum  presents, 
while  the  hypophysis  remains  within  the  sella  turcica.  Laterally,  the  tuber  cine- 
reum  is  bounded  by  the  tractus  optici,  whose  further  course  is  over  the  forward  and 
outwardly  coursing  cerebral  stalks,  the  pedunculi  cerebri,  and  then  to  pass  deeply. 
Behind  the  tuber  cinereum,  rise  two  white  pyriform  structures,  "the  corpora  mamil- 
laria  or  candicantia.  Behind  these  and  between  the  pedunculi  cerebri  lies  the  fossa 
interpeduncularis,  which  is  prolonged  backward  into  the  recessus  posterior  and  forward 
into  the  recessus  anterior.  The  floor  of  this  depression  is  formed  by  the  substantia 
perforata  posterior,  a  gray  surface  modelled  by  numerous  apertures  and  divided  into 
halves  by  a  median  furrow.  Towards  the  cerebral  peduncle  it  is  bounded  by  a 
groove,  the  sulcus  nervi  oculomotorii,  from  which  emerge  the  fibres  of  the  oculo- 
motor nerve. 

Behind  these  deeply  sunken  structures,  appears  a  white,  broad,  transverse  bridge, 
the  pons  Varolii,  which  in  front  and  behind  is  sharply  bounded,  in  the  middle  is  impressed 
by  a  broad  median  furrow,  the  sulcus  basilaris,  and  at  the  sides  narrows  and  then  extends 
laterally  and  backward  to  sink  into  the  cerebellum.  Behind  the  pons  lies  the  tapering 
bulb,  the  medulla  oblongata,  which  is  prolonged  into  the  spinal  cord.  It  presents  the 
median  longitudinal  furrow,  the  fissura  mediana  anterior,  that  is  bounded  on  each  side 
by  a  white  strand,  the  pyramid  or  pyramis.  Beyond  the  pyramidal  tract,  the  sulcus 
lateralis  anterior  extends  as  a  shallow  groove,  beyond  which,  in  turn,  lies  an  elongated 
egg-shaped  elevation,  the  oliva  or  olivary  eminence.  The  medulla  covers  the  median 
part  of  the  cerebellum,  occupying  a  broad  furrow,  known  as  the  vallecula  cerebelli, 
behind  which  appears  the  strongly  arched  ventral  surface  of  the  cerebellum.  A  deep 
median  cleft,  the  incisura  cerebelli  posterior,  separates  the  two  halves  of  the  little  brain,  the 
hemisphaeria  cerebelli,  which  exhibit  numerous,  more  or  less  parallel  narrow  tracts,  the 
folia.  On  slightly  raising  the  cerebellum,  the  fissura  transversa  cerebri  appears  as  a 
deep  cross  cleft,  separating  the  cerebellum  from  the  cerebrum  and  opening  into  the  fissura 
longitudinalis  cerebri. 


14  MORPHOLOGY. 

Closer  examination  of  the  base  of  the  brain  leads  further  to  the  location  of  the 
exits  of  the  individual  cerebral  nerves  from  the  brain,  concerning  which  the  following 
table  may  afford  explanation.  The  exits  of  these  nerves  from  the  skull  are  also  noted. 


Nerve 


I.  Fila  olfactoria 

II.  N.  opticus 

III.  N.  oculomotoris 

IV.  N.  troch'.earis 

V.  N.  trigeminus 

VI.  N.  abducens 

VII.  N.  facialis 

VIII.  N.  acusticus 

IX.  N.  glosso- 

pharyngeus 

X.  N.  vagus 

XI.  N.  accessorius 


XII.     N.  hypoglossus 


Exit  from  the  Brain 


Bulbus  olfactorius 
Chiasma  opticum 

Suicus  nervi  oculomotorii,  close  in  front 
of  pons,  on  medial  edge  of  cerebral 
peduncle 

Dorsal,  behind  the  corp.  quadrigemina, 
lateral  to  frenulum  veli  medullaris  an- 
terioris.  Course  around  the  cerebral 
peduncle 

Front  border  of  pons,  lateral,  near  the 
entrance  of  middle  cerebellar  peduncle 
into  the  cerebellum 

Hind  border  of  pons,  in  the  groove 
between  the  latter  and  the  medulla 
(pyramid) 

Lateral  to  N.  abducens,  on  hind  border 
of  pons.  in  front  of  and  lateral  to  olive 


Lateral  to  N.  facialis,  on  hind  border  of 
pons,  lateral  to  olive 

Behind  the  N.  facialis  and  N.  acusticus, 
in  upper  part  of  furrow  behind  olive 

Behind  the  N  glossopharyngeus,  in  the 
furrow  behind  the  olive 

Upper    root -fibres   (cerebral    portion): 
behind  N.  vagus,  in  the  furrow  behind 
•  the  olive 

Lower  root-fibres  (spinal  portion) :  be- 
tween the  front  and  hind  roots  of  the 
cervical  nerves,  as  far  as  sth  or  6th. 

Suicus  lateralis  anterior,  between  pyra- 
mid and  olive 


Exit  from  Skull 


Lamina  cribrosa 
Foramen  opticum 
Fissura  orbitalis  superior 


Fissura  orbitalis  superior 


R.  ophthalmicus:  Fis.  orbit,  sup. 
R.  maxillaris:  Foram.  rotundum 
R.  mandibularis:  Foram.  ovale 

Fissura  orbitalis  superior 


Porus  acusticus  internus— 
Meatus  acusticus  internus — 
Canalis  facialis 
Foramen  stylo-mastoideum 
Porus  acusticus 

Foramen  jugulare 
Foramen  jugulare 
Foramen  jugulare 


Canalis  hypoglossi 


N.  I,  II  and  VIII  are  sensory  nerves, 

N.  V,  VII,  IX  and  X  are  mixed  nerves, 

N.  III.  IV,  VI,  XI  and  XII  are  motor  nerves. 


Let  us  now  examine  a  median  sagittal  section  through  the  brain.  In  the  first 
place,  we  recognize  the  brain-mass  belonging  to  the  hemisphere,  with  its  fissures  and 
convolutions,  and,  further,  the  corpus  callosum,  the  large  commissure  connecting  the 


CORPUS  CALLOSUM.  15 

two  cerebral  hemispheres.  The  middle  part  of  the  bridge  is  the  body  (truncus  corporis 
callosi};  behind,  the  commissure  thickens  to  form  the  splenium ;  while  in  front,  it  bends 
sharply  downward  and  forms  the  knee,  genu  corporis  callosi,  that  tapers  into  the  beak- 
like  rostrum  corporis  callosi.  The  latter  is  prolonged  as  a  short  thin  white  lamella,  the 


Corpus  cnllosum   (Truncus) 

Sept.  pellncidnm 

Corpus  callosnm  (Genu) 


•idibulum 
Hypophysis     in- 

fundi-    Tub.    Corp. 
bnli  ciner.    mamilla 


Splenium  corp.  callis 
Corpus  pineale 

Corpora  quadrigemina 
Pedunculus  cerebri 


Medulla  otlongata 


FIG.   16. — Medis 


through  the  brain. 


FIG.  17. — Median  sagittal  section  through  the  adult  brain. 

lamina  rostralis,  which  is  continuous  with  the  attenuated  lamina  terminalis  that  extends 
to  the  front  surface  of  the  chiasma  opticum.  Behind  the  corpus  callosum,  covered  by 
the  hinder  part  of  the  hemisphere,  lies  the  cerebellum  ;  the  deep  fissura  transversa 
cerebri  is  plainly  seen  separating  the  hemisphere  and  cerebellum. 


i6 


MORPHOLOGY. 


Let  us  examine  the  parts  of  the  brain  lying  beneath  the  corpus  callosum.  Closely 
attached  to  the  under  surface  of  the  latter,  a  lamella  of  white  matter  extends  fonvard 
from  the  place  where  the  splenium  joins  the  body  or  trunk  of  the  corpus  callosum.  The 
structure  gradually  leaves  the  corpus  callosum,  arches  downward  with  forwardly  directed 
curve  until  close  behind  the  lamina  rostralis,  and  then  sinks  deeply  into  the  brain-sub- 
stance, just  behind  a  transversely  cut  white  bundle  of  fibres,  the  anterior  commissure  or 
commissura  anterior.  This  white  lamella  belongs  to  the  fornix.  Between  the  fornix, 
on  the  one  hand,  and  the  truncus,  genu,  rostrum  and  lamina  rostralis  of  the  corpus 
callosum,  on  the  other,  extends  a  thin  white  sheet,  the  septum  pellucidum.  Beneath  the 


Foramen  Monrot 


Massa  intermedia 


Gyms  cingnli 


Cyrus  snbcaltosns     —  ^ 


Area  parolfactoria. 
(Broca's  field) 


Corpus  mamillfire 
FIG.  18. — Median  sagittal  section  through  the  adult  brain;  subcallosal  region. 


fornix  and  the  hind  part  of  the  corpus  callosum  is  situated  the  thalamus,  between  whose 
fore-end  and  the  descending  fornix  lies  an  opening,  the  foramen  interventriculare  or  foramen 
of  Monro.  At  the  posterior  end  of  the  thalamus,  beneath  the  splenium  corporis  callosi, 
lies  the  pineal  body,  the  corpus  pineale.  The  cleft,  which  penetrates  the  pineal  body  in 
front,  is  called  the  recessiis  pinealis.  Immediately  beneath  is  found  the  cross-section  of 
the  commissura  posterior  cerebri,  with  which  are  joined,  proceeding  backward,  the  lamina 
quadrigemina,  the  velum  medullare  anterius  and  the  cerebellum.  On  the  median  surface 
of  the  thalamus,  behind  the  foramen  interventriculare,  lies  the  cross-section  of  the  middle 
commissure  or  massa  intermedia,  by  means  of  which  the  opposed  surfaces  of  the  two 
thalami  are  connected. 

The  sulcus  hypothalamicus   (Monroi)    is  a  furrow  that  extends  backward  from   the 
foramen  interventriculare,  beneath  the  massa  intermedia,  towards  the  commissura  posterior 


TELENCEPHALON.  17 

and  separates  the  region  of  the  thalamus  from  the  more  dependent  hypothalamus.  On 
examining  this  region  more  closely,  we  note  again  parts  that  have  been  mentioned  in 
connection  with  the  base  of  the  brain :  in  front  the  lamina  terminalis  that  joins  the 
anterior  surface  of  the  chiasma  opticum,  the  recessus  options  between  the  lamina  and  the 
chiasma  and  behind  the  latter,  the  recessus  infundibuli,  the  infundibulum  with  the  hypoph- 
ysis, the  tuber  cinereum,  the  corpus  mamillare,  and  the  substantia  perforata  posterior, 
forming  the  floor  of  the  fossa  interpeduncidaris  (  Tarini). 

Continuing  backward,  the  cerebral  peduncle,  the  pons  and  the  medulla  oblongata 
are  seen  in  cross-section.  The  sulcus  hypothalamicus,  running  backward  from  the 
foramen  interventriculare,  opens  into  the  aquaedudus  cerebri,  or  aqueduct  of  Sylvius, 
which  extends  beneath  the  quadrigeminal  plate  and  joins  the  fourth  ventricle  that  under- 
lies the  cerebellum  (Figs.  16,  17,  and  18). 

TELENCEPHALON. 

The  telencephalon  or  end-brain  includes  : 
The  hemisphaerium, 
The  pars  optica  hypothalami. 
To  the  hemisphaerium  belong : 

The  pallium  or  cerebral  mantle, 
xThe  rhinencephalon, 

The  stem  of  the  telencephalon — the  gray  nuclei  of  the  end-brain. 
To  the  pars  optica  hypothalami  belong  : 
The  lamina  terminalis, 
The  chiasma  opticum, 
The  tuber  cinereum, 
The  infundibulum, 
The  hypophysis. 
The  hemisphaerium  contributes  the  chief  mass  of  the  end-brain. 

In  order  to  study  the  morphology  of  the  telencephalon  to  the  best  advantage,  one 
proceeds  in  the  following  manner :  the  brain  is  placed  on  the  dorsal  surface,  with  the 
base  upward ;  the  pons,  cerebellum  and  medulla  oblongata,  all  connected,  are  completely 
separated  from  the  brain  by  a  transverse  cut  passing  through  the  front  border  of  the 
pons.  A  second  cut,  sagittal  and  in  the  mid-line,  divides  the  two  hemispheres  from 
each  other. 

In  the  first  place,  let  us  examine  a  hemisphere  in  general.  Each  hemisphere 
presents  three  surfaces  :  a  convexly  arched  dorso-lateral  surface,  a  flat  median  surface, 
and  a  basal  surface,  which  is  subdivided  by  a  deep  incision  into  a  smaller  anterior  and 
a  larger  posterior  part.  We  distinguish  further  an  anterior  frontal  pole  (^polus  frontalis}, 
a  posterior  pole  {polus  occipitalis)  and  a  temporal  pole  {polus  temporalis} ,  the  latter 
representing  the  fore-end  of  the  posterior  division  of  the  basal  surface.  A  dorsal  border 
marks  the  transition  of  the  lateral  to  the  median  surface  ;  its  medial  continuation  forms 
the  base  to  the  basal  border.  The  lateral  border  corresponds  to  the  transition  of  the 
lateral  at  the  basal  surface. 


I8  MORPHOLOGY. 


PALLIUM— CEREBRAL  MANTLE. 

The  surface  of  the  pallium  or  cerebral  mantle  is  subdivided  into  definite  lobes  (lobi) 
by  definite  and  usually  deep  clefts  and  furrows,  the  fissures  and  sulci.  Of  such  divisions 
or  lobi  cerebri  are  recognized  : — 

Lobus  frontalis, 

Lobus  parietalis, 

Lobus  temporalis, 

Lobus  occipitalis. 

An  additional  special  lobe,  the  insula  or  island  of  Reil,  lies  hidden  at  the  bottom 
of  the  lateral  or  Sylvian  fissure.  Each  lobe  further  exhibits  convolutions  (gyri  cerebri}, 
which,  while  bounded  by  the  fissures,  are  often  connected  at  the  bottom  of  the  fissures 

Sulc    frontal.      Sulc.  front.  'Sulc.  Jrae-     Sulc.  centralis     Sulc.  post-  , 

ed.  *«/.  central,  sup.         (Rolandi)  centralis    Sulc.  interpariet. 

~*  .  /  transfers. 

Sulc.  praecentral.  ^ 

Sulc.  frontal,  inf. 

Ram.  ant.  ascend.  - Sulci  occipital 

Ram.  ant.  hori-     -    '^  --  ***erioret  et  later- 

Truncus  fUsurae 

Fitsura  cerebri    Polui  tern-     Sulc.  temp.        Sulc.        Sulc.  Incisura  praeoccipital. 

lateral.  (Ram.  post.)       for  alls  sup.          temp.  med.  temp.  inf. 

FIG.  19. — Dorso-lateral  cerebral  surface.    Fissures  and  convolutions. 

by  deep  convolutions  (gyri  profundi).  The  short  superficial  or  sunken  convolutions 
that  connect  two  longer  gyri  are  called  annectant  convolutions  (gyri  transitivi} .  The 
secondary  fissures  (incisura)  are  superficial  aberrant  furrows,  usually  uncertain  in  their 
course  and  springing  from  deeper  sulci,  that  cut  into  the  convolutions  and  in  certain 
cases  cause  doubling  of  the  gyri. 

LOBES  AND  GYRI  OF  THE  DORSO-LATERAL  SURFACE. 

Turning  again  to  the  basal  aspect  of  the  hemisphere,  the  vallecula  Sylvii  (fossa 
cerebri  lateralis)  appears  as  a  deep  cleft,  lateral  to  the  substantia  perforata  anterior,  that 
separates  the  basis  cerebri  into  an  anterior  and  posterior  division.  From  the  valley  the 
fissura  cerebri  lateralis,  or  Sylvian  Jissure,  extends  outward,  at  first  as  the  truncus  fissurae 
lateralis,  toward  the  dorso-lateral  surface  of  the  hemisphere.  On  reaching  the  latter,  the 
fissure  divides  into  three  branches  :  ( i )  the  short  ramus  anterior  horizontalis ;  running 
horizontally  forward,  (2)  the  ramus  anterior  ascendens,  also  short  and  directed  almost 
vertically  upward,  and  (3)  the  long  ramus  posterior,  which  continues  the  direction  of  the 


FISSURES   AND   CONVOLUTIONS.  19 

anterior  horizontal  limb  backward  and  somewhat  obliquely  upward  and  at  its  end  usually 
divides  in  a  Y-like  manner  into  a  ramus  ascendens  and  a  ramus  descendens.  Approxi- 
mately from  the  middle  of  the  dorsal  border  of  the  hemisphere,  the  sulcus  centralis  or 
fissure  of  Roland,  runs  obliquely  downward  and  forward  toward  the  posterior  ramus  of 
the  fissura  cerebri  lateralis.  As  a  rule,  this  furrow  exhibits  two  knee-like  bends,  one  at 
the  junction  of  the  upper  and  middle  thirds,  the  other  at  the  transition  of  the  second 
and  lower  thirds  ;  the  fissure,  moreover,  usually  crosses  the  upper  border  of  the 
hemisphere. 

Lobus  frontalis.  The  frontal  lobe  lies  above  the  fissure  cerebri  lateralis  and  in 
front  of  the  central  fissure,  and  presents  the  following  fissures  and  convolutions.  The 
sulcus  praecentralis  superior  begins  somewhat  below  the  upper  border  of  the  hemisphere 
and  runs  more  or  less  parallel  with  the  sulcus  centralis.  Somewhat  lower,  the  sulcus 
praecentralis  inferior  continues  in  the  same  direction  and  below  penetrates  between  the 
ramus  anterior  ascendens  fissurae  cerebri  lateralis  and  the  lower  end  of  the  sulcus  cen- 
tralis. The  upper  end  of  the  inferior  precentral  sulcus  almost  constantly  lies  in  advance 
of  the  lower  end  of  the  superior  fissure.  As  variations,  the  precentral  fissures  may  con- 
nect with  the  central  fissure  and  the  lower  precentral  may  join  the  fissura  cerebri  lateralis. 

The  sulcus  frontalis  superior  extends  forward  from  the  superior  precentral  sulcus, 
approaching  the  upper  border  of  the  hemisphere  in  front.  At  times  the  fissure  cuts 
through  the  sulcus  praecentralis  superior  towards  the  central  fissure,  thereby  producing 
the  cruciform  type  of  precentral  furrow.  In  many  cases  the  superior  frontal  sulcus  is 
interrupted  by  two  or  three  annectant  convolutions.  The  fissure  may  also  be  doubled. 

The  sulcus  frontalis  inferior  likewise  extends  forward,  from  the  inferior  precentral 
fissure,  but  more  arched  and  downward.  The  fissure  is  usually  clearly  marked,  but  it 'may 
present  very  variable  forms  and  be  interrupted  by  deep  or  superficial  annectant  convolutions. 
Ordinarily  a  short  furrow,  the  sulcus  radiatus,  extends  downward  from  the  inferior 
frontal  between  the  anterior  horizontal  and  ascending  rami  of  the  fissura  cerebri  lateralis. 

The  small  sulcus  frontalis  medius  is  generally  to  be  seen  between  the  superior 
and  inferior  frontal  fissures.  This  sulcus  is  often  readily  identified,  but  it  may  exhibit 
the  most  diverse  forms,  since  it  may  be  displaced  or  effaced  by  annectant  convolutions. 
At  times  the  fissure  is  clearly  recognizable  as  a  continuous  and  deep  furrow. 

The  foregoing  fissures  bound  the  following  convolutions.  The  gyrus  centralis 
anterior  lies  between  the  superior  and  inferior  precental  fissures  in  front  and  the  central 
fissure  behind.  The  gyrus  frontalis  superior  is  bounded  by  the  superior  frontal  fissure 
below  and  the  superior  prefrontal  fissure  behind.  Between  the  superior  and  inferior 
frontal  fissures  extends  the  gyrus  frontalis  medius,  which  is  subdivided  by  the  median 
frontal  sulcus  into  a  pars  superior  and  a  pars  inferior.  The  gyrus  frontalis  inferior  lies 
below  the  inferior  frontal  fissure.  This  convolution,  also  known  as  Brocd s  convolution 
on  the  left  side,  includes  three  subdivisions: — 

The  pars  opercidaris ,  between  the  lower  end  of  the  inferior  precentral  fissure  and 
the  anterior  ascending  ramus  of  the  fissura  cerebri  lateralis; 

The  pars  triangularis ,  between  the  anterior  ascending  and  horizontal  rami  of  the 
lateral  fissure  ;  and 

The  pars  orbitalis,  between  the  anterior  horizontal  ramus  and  the  trunk  of  the 
lateral  fissure. 


20  MORPHOLOGY. 

Behind  the  sulcus  centralis,  or  fissure  of  Rol.mdo,  and  above  the  ramus  posterior 
of  the  fissure  of  Sylvius,  stretches  the  parietal  lobe,  while  below  the  last  named  fissure 
lies  the  temporal  lobe.  Posteriorly,  both  lobes  pass  into  the  occipital  lobe  without  a 
definite  boundary.  As  a  conventional  boundary,  we  may  adopt  a  line  that  unites  the 
dorsal  end  of  the  parieto-occipital  fissure,  which  incises  the  upper  border  of  the  hemi- 
sphere, with  the  incisura  praeoccipitalis.  The  fissura  parieto-occipitalis  is  a  deep  cleft  on 
the  hind  part  of  the  median  surface  of  the  hemisphere  (Fig.  22),  which  incises  the  upper 
border  of  the  hemisphere  and  extends  a  short  distance  on  its  dorso-lateral  aspect.  It 
is  readily  identified  as  a  deep  incision  on  the  upper  border  of  the  hemisphere  about 
midway  between  the  central  fissure  and  the  occipital  pole,  rather  nearer  the  latter.  The 
incisura  praeoccipitalis  appears  as  a  slight  notch  on  the  lateral  border  of  the  hemisphere, 
approximately  at  the  junction  of  the  middle  and  posterior  thirds  (Fig.  19). 

Lobus  parietalis.  The  sulcus  postcentralis  extends  behind  and  more  or  less 
parallel  with  the  central  fissure.  This  furrow  is  sometimes  continuous,  and  sometimes 


Gyr.  frontalis  superior      Gyr.  centralis  anterior      Gyr.  centr.  posterior        Lobulus  parietalii 


Gyr.  temporalis      Gyr.  temp. 
superior  medius 


Gyr.  descendens  (  Ecker  \ 


Gyr.  temporalis  inferior 
FIG.  20. — Dorso-lateral  cerebral  surface.     Fissures  and  convolutions. 


subdivided  into  two  parts,  the  sulcus  postcentralis  superior  and  inferior.  Each  sub- 
division may  retain  its  individuality,  or  at  the  same  time  join  the  sulcus  interparietalis. 
When  the  superior  postcentral  fissure  is  independent,  it  usually  exhibits  variations  in 
form  and  size,  sometimes  being  unbranched  and  paralleling  the  central  fissure,  but  often 
forming  a  three-  or  four-limbed  furrow.  As  does  the  precentral,  so  the  postcentral  fissure 
at  times  anastomoses  with  the  central  fissure ;  the  inferior  postcentral  fissure,  moreover, 
may  connect  with  the  Sylvian  fissure. 

The  sulcus  interparietalis  begins,  mostly  with  a  bifurcation,  behind  the  upper  end 
of  the  inferior  postparietal  fissure.  By  the  junction  of  this  sulcus  with  one  or  other  of 
the  postcentral  fissures,  a  veritable  vortex  of  furrows,  a  fissure-star,  is  formed.  The 
sulcus  interparietalis  pursues  an  arched  course  backward,  beneath  the  dorso-lateral  end  of 
the  parieto-occipital  fissure,  and  usually  opens  out  into  the  sulcus  occipitalis  transversus. 
Occasionally  the  interparietal  fissure  passes  across  the  transverse  occipital  furrow  and 
continues  backward  as  the  sulcus  occipitalis  superior.  The  interparietal  fissure  is  often 
made  up  of  several  parts  ;  during  its  course  isolated  fissures  are  given  off  upward  as 


FISSURES   AND   CONVOLUTIONS.  21 

well  as  downward.  A  short  furrow,  known  as  the  sulcus  parietalis.  transversus  (Bris- 
saud),  extends  upward,  towards  the  border  of  the  hemisphere,  from  in  front  of  the 
dorsal  end  of  the  parieto-occipital  fissure.  Often  two  furrows  pass  downward.  One 
sulcus  runs  behind  the  ascending  end-branch  of  the  ramus  posterior  of  the  Sylvian 
fissure  and  is  called  the  sulcus  intermedius  primus  (Jensen).  It  often  extends  as  a 
continuation  of  the  upper  transverse  parietal  fissure,  but  may  be  strongly  developed  and, 
indeed,  may  establish  a  connection  between  the  interparietal  fissure  and  the  ascending 
end  of  the  superior  temporal  fissure.  The  other  sulcus  is  given  off  farther  backward, 
runs  behind  the  ascending  end  of  the  superior  temporal  fissure  and  is  known'  as  the 
sulcus  intermedius  secundus  (Eberstaller).  Both  of  these  sulci  intermedii  may  also  exist 
as  independent  fissures. 

By  means  of  the  foregoing  fissures  the  following  convolutions  are  defined  :  the 
gyrus  centralis  posterior  lies  behind  the  sulcus  centralis,  bounded  below  by  the  fissura 
cerebri  lateralis  and  behind  by  the  fissura  postcentralis.  Above  the  sulcus  interpari- 
etalis  lies  the  lobulus  parietalis  superior,  while  beneath  this  fissure  extends  the  lobulus 
parietalis  inferior.  This  lower  parietal  lobule  presents  two  special  convolutions,  the 
gyrus  supramarginalis  and  the  gyrus  angularis.  The  gyrus  supramarginalis  encloses  the 
ascending  terminal  stem  of  the  ramus  posterior  of  the  Sylvian  fissure  and  is  bounded  by 
the  sulcus  intermedius  primus  behind.  The  gyrus  angularis  surrounds  the  ascending 
end  of  the  superior  temporal  fissure,  and  is  bounded  in  front  by  the  sulcus  intermedius 
primus  and  behind  by  the  sulcus  intermedius  secundus. 

Lobus  temporalis.  One  of  the  most  constant  fissures  is  the  sulcus  temporalis 
superior.  It  begins  in  front  at  the  temporal  pole,  extends  backward  and  upward  parallel 
to  the  fissura  cerebri  lateralis  and  ends,  as  a  rule,  in  the  gyrus  angularis  by  running 
upward  behind  the  ascending  terminal  branch  of  the  fissura  cerebri  .lateralis.  At  times 
one  finds  a  forking  into  an  ascending  and  a  descending  branch.  The  sulcus  temporalis 
medius  runs  below  the  superior  temporal  fissure.  The  fissure  is  seldom  continuous, 
usually  being  made  up  of  several  parts.  Below  the  middle  temporal  fissure,  and  on  the 
basal  surface,  extends  the  sulcus  temporalis  inferior.  The  three  temporal  convolutions 
are  defined  by  these  fissures.  The  gyrus  temporalis  siiperior  extends  below  the  sulcus 
cerebri  lateralis  and  above  the  superior  temporal  fissure  ;  the  gyrus  temporalis  medius 
lies  between  the  superior  and  middle  temporal  sulci ;  and  the  gyrus  temporalis  inferior 
is  located  below  the  inferior  temporal  fissure.  The  surface  of  the  upper  temporal  con- 
volution facing  the  Sylvian  fissure  presents  the 
gyri  temporales  transversi,  also  known  as 
the  convolutions  of  Heschl,  which  are  weakly 
developed  in  the  front  half  and  more  strongly 
behind. 

Lobus  occipitalis.     The  anterior  bound- 

Sulc.  centralis 

ary  of   the   lobus  occipitalis  is  formed  in  part          FlG  2I._Fissures  and  convolutions  of  the  insula. 
by  the  sulcus    occipitalis  transversus,  a    sulcus 

liable  to  many  variations  respecting  its  position,  length  and  direction.  In  addition,  there  are 
the  sulci  occipitales  superiores  and  the  sulci  occipitales  laterales.  By  means  of  these  fissures 
the  gyri  occipitales  superiores  and  the  gyri  occipitales  laterales  are  defined.  Towards  the  oc- 
cipital pole,  the  convolutions  join  a  vertical  gyrus,  known  as  the  gyrus  descendens  (Ecker). 


22 


MORPHOLOGY. 


Insula.  On  penetrating  the  depth  of  the  fissura  cerebri  lateralis,  by  drawing 
apart  the  edges  of  the  bounding  lobes,  one  comes  to  a  deep  depression,  the  fossa 
cerebri  lateralis  (^Sylvii),  at  the  bottom  of  which  lies  the  insula,  also  known  as  the  basic 
lobe  (Stammlappen).  Those  parts  of  the  lobes  bounding  the  Sylvian  fissure,  which  cover- 
in  the  island,  together  constitute  the  operculum.  Since  the  frontal,  parietal  and  temporal 
lobes  participate  in  its  production,  we  distinguish  a  pars  frontalis,  a  pars  parietalis  and 
a  pars  temporalis  of  the  operculum.  The  surface  of  the  temporal  lobe  directed  towards 
the  insula  presents  the  sulci  and  gyri  temporales  transversi.  Similar  fissures  and  con- 
volutions exist  also  upon  the  surfaces  of  the  parietal  and  frontal  opercula  facing  the 
island.  The  insula  appears  in  the  form  of  an  irregular  conical  projection,  a  three-sided 
pyramid  whose  apex — the  island-pole — is  directed  forward  and  outward.  The  island  is 


Corpus  caHosum 


Sulcus  cinguli 
Gyr  frontal  sup 


Sulc   supraorbital. 
Sulc  parolfactor. 


Lobulus 
Snlcns  paracentral'a      paracentr.          Sulcus  centralis 


Ramus  marginal. 
Sulci  cingnli 


Sulcus  temporalis  inf. 
Fissura.  rhinica    Fissufa.  hippocampi        Gyr.  fusiformis 
FIG.  22. — Medial  cerebral  surface.     Fissures  and  convolutions. 


encircled  by  a  deep  fissure,  the  sulcus  circularis  (Reili).  Since  this  furrow,  strictly 
considered,  is  not  circular  but  rather  triangular,  a  sulcus  anterior,  a  sulcus  superior  and 
a  sulcus  inferior  may  be  distinguished.  The  sulcus  anterior  separates  the  island  from 
the  orbital  part  of  the  frontal  operculum,  the  sulcus  superior  from  the  fronto-parietal 
operculum,  and  the  sulcus  inferior  from  the  temporal  operculum.  The  island  is  divided 
into  a  lobus  insulae  anterior  and  a  lobus  insulae  posterior  by  the  sulcus  centralis  insulae, 
a  fissure  that  runs  from  in  front  and  below  backward  and  upward.  The  anterior  lobule 
exhibits  several  short  convolutions,  the  gyri  breves  insulae,  while  the  posterior  lobule 
appears  as  the  gyrus  longus  insulae,  which  now  and  then  is  subdivided  into  two  con- 
volutions by  a  long  furrow  which  parallels  the  sulcus  centralis  insulae. 


LOBES  AND  GYRI  OF  THE  MEDIAL  AND  BASAL  SURFACES. 

All  four  cerebral  lobes,  with  which  we  have  now  become  somewhat  intimately 
acquainted  on  the  dorso-lateral  aspect  of  the  hemisphere,  are  continued  onto  the  medial 
and  partly  also  onto  the  basal  surface.  They  do  not  extend,  however,  over  the  entire 
medial  surface,  but  bound  a  large  annular  tract  that  belongs  to  the  rhinencephalon. 
Let  us  first  examine  the  defining  fissures  and  sulci. 


FISSURES   AND   CONVOLUTIONS. 


The  sulcus  cinguli,  or  calloso-marginal  fissure,  begins  beneath  the  rostrum  of  the 
corpus  callosum.  It  runs  forward,  around  the  knee,  and  then  backward,  more  or  less 
parallel  with  the  corpus  callosum,  as  far  as  the  splenium.  Here  it  bends  at  a  blunt 


'Sulc-us  parolfact. 
Post. 


Sulcns  parolfact. 


Isthmus 
Vssura  rkk  Gyr.  hippoc. 

FIG.  23. — Medial  cerebral  surface.      Gyrus  fornicatus  is  shaded. 

angle  upward  towards  the  superior  margin  of  the  hemisphere  as  the  ramus  marginalis. 
During  its  entire  course,  several  and  sometimes  deep  incisions  branch  off,  upward  as 
well  as  downward.  In  front  of  the  obtuse  bend,  approximately  over  the  middle  of  the 

Sulcns  olfactorius 


Pissura  collateral. 


Fissura  calcarina 


FIG.  24. — Basal  surface  of  the  brain.     Fissures  and  convolutions. 

corpus  callosum,  the  fissure  usually  sends  a  side  branch,  the  sulcus  paracentralis,  upward. 
Another  branch,  the  sulcus  supraorbitalis  (Broca),  is  given  off  at  the  level  of  the  genu. 
Finally,  a  third  fissure,  the  sulcus  subparietalis ,  which  represents  a  continuation  of  the 
chief  furrow,  runs  backward  and  around  the  splenium  corporis  callosi.  Immediately 
beneath  the  knee  and  the  rostrum  of  the  callosum,  the  sulcus  corporis  callosi  begins,  at 


24  MORPHOLOGY. 

first  as  only  a  shallow  fissure.  It  often  appears  there  as  the  prolongation  of  the  sulcus 
parolfactorius  posterior  (see  Rhinencephalon,  page  26),  then  continues  around  the  genu, 
closely  follows  the  convex  surface  of  the  corpus  callosum,  runs  around  the  splenium 
and  continues  into  the  fissura  hippocampi,  the  deep  cleft  that  runs  from  behind  and 
above  forward  and  downward. 

In  the  posterior  part  of  the  medial  surface  of  the  hemisphere,  beginning  about  mid- 
way between  the  turned-over  end  of  the  central  fissure  and  the  occipital  pole,  the  deep 
fissura  parieto-occipitalis  runs  obliquely  forward  and  downward,  behind  the  lower  end 
of  the  subparietal  branch  of  the  sulcus  cinguli,  as  far  as  the  region  beneath  the  splenium 
corporis  callosi.  In  the  lower  part,  at  about  the  level  of  the  splenium,  the  furrow  is 
joined  at  an  acute  angle  by  the  deep  fissura  calcarina.  The  latter,  slightly  arched 
and  somewhat  above  the  medial  border,  extends  backward  toward  the  occipital  pole, 
where  it  may  end  as  a  simple  groove,  or,  as  is  usually  the  case,  in  two  widely  divergent 
branches.  Occasionally  the  calcarine  fissure  overruns  the  occipital  pole  and  terminates 
on  the  dorso-lateral  surface  of  the  hemisphere.  The  stem  formed  by  the  union  of  the 
parieto-occipital  and  calcarine  fissures  extends  downward  and  close  behind  the  hippo- 
cam  pal  fissure,  without,  however,  joining  the  latter.  The  fissura  collateralis  begins  at 
the  level  of  the  occipital  pole,  below  the  calcarine  fissure,  and  passes  forward  below  the 
common  stem  of  the  parieto-occipital  and  calcarine  fissures.  Its  continuation  into  the 
anterior  part  of  the  temporal  lobe  constitutes  the  fissura  rhinica,  whose  front  end  is 
known  as  the  incisura  temporalis  (Schwalbe).  Below  the  collateral  fissure  is  the  sulciis 
temporalis  inferior. 

By  means  of  the  foregoing  fissures,  the  following  parts  are  defined.  The  tract 
occupying  the  front  part  of  the  medial  surface  outside  the  sulcus  cinguli  belongs  to  the 
frontal  lobe,  more  particularly  to  the  superior  frontal  convolution.  It  extends  back- 
ward beyond  the  paracentral  sulcus,  its  posterior  limit  being  a  line  drawn  from  the 
medial  end  of  the  central  sulcus,  between  the  paracentral  and  marginal  rami  of  the 
sulcus  cinguli.  to '  the  last-named  fissure.  The  tract  between  the  paracentral  and  mar- 
ginal branches  of  the  sulcus  cinguli  is  called  the  lobuhis  paracentralis.  Here  is  found  the 
transition  of  the  gyrus  centralis  anterior  into  the  gyrus  centralis  posterior.  The  larger 
part  of  the  paracentral  lobule  belongs  to  the  precentraK  convolution.  Behind  the  tract 
belonging  to  the  frontal  lobe,  a  region  broadens  out  which  belongs  to  the  parietal  lobe. 
It  Kes  above  the  sulcus  cinguli  and  its  prolongation,  the  subparietal  fissure,  and  is 
bounded  behind  by  the  parieto-occipital  fissure.  The  entire  tract,  between  the  marginal 
arm  of  the  sulcus  cinguli  in  front,  the  subparietal  fissure  below  and  the  parieto-occipital 
fissure  behind,  constitutes  the  praecuneus  or  quadrate  lobule.  Between  the  parieto-occip- 
ital and  calcarine  fissures  lies  the  cuneus,  which  belongs  to  the  occipital  lobe.  Below 
the  calcarine  fissure,  between  it  and  the  collateral  fissure,  lies  another  part  of  the  occip- 
ital lobe  known  as  the  gyrus  lingualis.  On  the  basal  aspect  of  the  hemisphere,  below 
the  collateral  fissure,  the  gyrus  fusiformis  extends  as  a  part  of  the  temporal  lobe.  It 
is  also  called  the  gyrus  occipito-temporalis. 

A  large  annular  tract  belonging  to  the  rhinencephalon  is  enclosed  by  the  foregoing 
convolutions  and  lobes.  Externally  it  is  bounded  by  the  sulcus  cinguli,  the  common 
stem  of  the  parieto-occipital  and  calcarine  fissures,  the  front  end  of  the  collateral  fissure 
and  the  fissura  rhinica.  The  inner  boundary  is  contributed  by  the  sulcus  corporis  callosi 


RHINENCEPHALON.  25 

and  the  fissura  hippocampi.  In  its  entirety,  this  tract  constitutes  the  gyrus  fornicatus 
or  the  limbic  lobe.  It  is  subdivided  into  the  gyrus  cinguli,  which  arches  around  the 
corpus  callosum,  and  the  gyrus  hippocampi,  which  is  included  between  the  hippocampal 
fissure  on  the  one  side  and  the  collateral  and  rhinal  fissures  on  the  other  and  hooks 
around  the  anterior  end  of  the  hippocampal  fissure  to  form  the  uncus.  The  gyrus 
cinguli  and  the  gyrus  hippocampi  are  continuous,  behind  and  beiow  the  splenium,  by 
means  of  the  isthmus  gyri fornicati. 

Turning  once  more  to  the  basal  surface,  in  the  posterior  and  larger  division,  we 
note  the  sulci  and  gyri  already  mentioned: — the  fissura  hippocampi,  the  fissura  parieto- 
occipitalis  and  calcarina,  joining  to  form  a  common  stem,  the  fissura  collateralis,  the 
fissura  rhinica,  the  sulcus  temporalis  inferior  and  the  convolutions  extending  between 
these  furrows.  The  anterior  and  smaller  division  of  the  basal  aspect  belongs  to  the 
lobus  frontalis,  being  known  as  its  orbital  surface.  Near  the  medial  border  of  the  latter, 
the  straight  sulcus  olfactorius  runs  forwards  and  somewhat  medially  and  lodges  the 
olfactory  bulb  and  tract.  The  sulcus  is  deep  and  almost  always  extends  farther  forwards 
than  the  anterior  end  of  the  bulbus  olfactorius.  Behind,  it  divides  into  a  ramus  medialis 
and  lateralis,  which  embrace  the  tuberculum  olfactorium.  Lateral  to  the  olfactory  fissure 
lie  some  furrows  of  uncertain  number  and  arrangement,  the  sulci  orbitales.  By  their  union 
the  most  varying  patterns  are  produced,  including  H-,  X-,L-,T-,K-  and  Z  - 
like  forms.  Medial  from  the  sulcus  olfactorius  extends  the  gyrus  rectus.  The  gyri 
orbitales  are  bounded  by  the  orbital  fissures. 

RHINENCEPHALON. 

The  rhinencephalon  embraces  :  (a)  the  peripheral  division  and  (£)  the  central  or 
cortical  division. 

The  peripheral  division  includes  the  lobus  olfactorius,   to  which  belong: 
The  bulbus  olfactorius, 
The  tractus  olfactorius, 
The  tuberculum  olfactorium,  with  the  gyri 

olfactorii  medialis  and  lateralis, 
The  area  parolfactoria  (Broca), 
The  substantia  perforata  anterior, 
The  gyrus  diagonalis  rhinencephali, 
The  gyrus  subcallosus  (Zuckerkandl}. 

The  central  or  cortical  division  includes  : 

The  gyrus  fornicatus  (Arnold}, 

The  hippocampus, 

The  gyrus  dentatus, 

The  gyrus  uncinatus, 

The  gyrus  intralimbicus, 

The  gyrus  fasciolaris, 

The  gyri  Andreae  Retzii  or  callosal  gyri. 


26 


MORPHOLOGY. 


i.  LOBUS   OLFACTORIUS. 

The  olfactory  lobe  falls  under  two  subdivisions,  one  in  front,  the  lobus  olfactorius 
anterior,  and  one  behind,  the  lobus 
olfadorius  posterior  (Figs.  25  and 
26).  These  are  separated  from 
each  other  by  a  fissure,  the  sul- 
cus  parolfactorius  posterior  (the 
embryonic  fissura  prima  of  His), 
which  runs  behind  the  trigonum 
olfactorium,  between  the  latter  and 
the  substantia  perforata  anterior, 
and  continues  toward  the  medial 
surface  of  the  hemisphere.  A 

To   the    lobus    olfactorius 
anterior  belong : 

The    bulbus    olfactorius, 
The   tractus    olfactorius, 

The  tuberculum  olfac- 
torium and  the  di- 
verging gyri  olfactorii 
medialis  and  lateralis. 


The  area  parolfactoria. 


Trigonum 
Gyrns  olfact.  lateralis 

Gyrus  olfactorio-orbitalis 
Subst.  Perf.  ant. 
Insula 


Angvlus  gyr    olf.  lateral. 


Gyr.  ambiens 
Sulc.  inf.  rhinenceph. 
Gyr.  semilunar 
Sulcus  semiannularis 


FIG.  25. — Brain  of  human  foetus  of  between  five  and  six  months. 
Basal  aspect. 


Sulcus  parolfact.  post. 


Gyr.  olfactorius  medialis 
Br oca's  diagon.  band 


Trigon. 

praecommissurale 
(Lamina  terminala) 


FIG.  26. — Schematic  representation  of  the  lobus  olfactorius. 


LOBUS   OLFACTORIUS   ANTERIOR.  27 

To  the  lobus  olfactorius  posterior  belong: 

The  subtiantia  perforata  anterior  or  gyrus  perforatus  rhinencephali, 
The  diagonal  band  of  Broca,   or  gyrus  diagonalis  rhinencephali, 
The  gyrus  subcallosus. 

A.     LOBUS  OLFACTORIUS  ANTERIOR. 

The  bulbus  olfactorius  presents  usually  an  oval  form — an  ellipse  or  a  vertically 
flattened  band — and  constitutes  an  enlargement  of  the  tractus  olfactorius  in  front.  On 
the  under  surface,  delicate  threads,  the  fila  olfactoria,  pass  out  and  descend  into  the 
nasal  fossa  through  the  apertures  of  the  lamina  cribrosa.  They  are  disposed  in  two 
series  and  may  be  designated  as  the  fila  olfactoria  medialia  and  lateralia.  They  are  so 
delicate  that  they  are  always  broken  off  when  the  brain  is  removed. 

The  tractus  olfactorius  lies  as  a  white  strand  in  the  olfactory  sulcus  and, 
on  cross-section,  reveals  the  form  of  a  triangle  with  base  below  and  apex  above  buried 
in  the  sulcus.  The  hind  part  of  the  tract,  toward  the  tuberculum  olfactorium,  is  narrow 
and  somewhat  compressed. 

The  tuberculum  olfactorium,  into  which  the  tractus  is  prolonged  behind, 
appears  in  its  true  form  only  after  the  bulb  and  tract  have  been  raised  from  the 
olfactory  sulcus  and  the  latter  itself  rendered  more  gaping  by  pulling  apart  the  bounding 
convolutions.  Then  the  tuberculum  appears  as  a  pyramidal  elevation,  whose  apex  pene- 
trates the  sulcus  and  whose  base  forms  an  irregular  triangular  field,  the  trigonum 
olfactorium. 

From  the  tuberculum  proceed  the  medial  and  lateral  olfactory  convolutions  whose 
courses  are  as  follows  : — 

The  gyrus  olfactorius  medialis  runs  as  a  narrow  convolution  medially.  In  front, 
it  is  bounded  by  the  medial  posterior  branch  of  the  sulcus  olfactorius;  medially  and 
behind,  by  the  sulcus  parolfactorius  posterior  (the  fissura  prima  of  His).  A  white  fascic- 
ulus of  fibres,  the  stria  olfactoria  medialis,  the  continuation  of  the  medial  strand  of  the 
olfactory  tract,  streams  into  the  gyrus  olfactorius  medialis,  soon  to  become  lost  in  the 
gray  substance  of  the  convolution. 

On  following  the  medial  gyrus  farther,  it  is  found  to  radiate  within  a  small  field 
on  the  medial  surface  of  the  hemisphere,  that  lies  immediately  below  the  rostrum  of  the 
corpus  callosum  and  is  bounded  both  in  front  and  behind  by  a  small  fissure.  The  furrow 
in  front  is  the  sulcus  parolfactorius  anterior,  while  the  one  behind  is  the  continuation 
of  the  sulcus  parolfactorius  posterior,  already  mentioned.  The  small  field  is  called  the  area 
par  olfactoria,  or  field  of  Broca.  It  joins  the  gyrus  olfactorius  medialis  with  the  gyrus 
fornicatus,  particularly  with  the  gyrus  cinguli  (Figs.  18  and  23),  and  thus  establishes  the 
connection  of  the  lobus  olfactorius  anterior  with  the  central  region  of  the  rhinencephalon. 

The  gyrus  olfactorius  lateralis  runs  laterally.  In  the  foetal  brain  of  from  four 
to  five  months,  one  readily  recognizes  that  the  gyrus  passes  outward  towards  the  Sylvian 
fossa,  its  so-called  front  limb  proceeding  from  the  trigonum  almost  at  right  angles ; 
then,  at  the  medial  margin  of  the  fossa  and  after  an  acute  bend,  the  hind  limb  runs 
backward  and  medially  to  the  anterior  border  of  the  gyrus  hippocampi.  Here  the  gyrus 
ends,  to  a  certain  extent,  in  two  claws,  the  medial  one  of  which  is  known  as  the  gyrus 
semihinaris  rhinencephali,  and  the  lateral  as  the  gyrus  ambiens  rhinencephali.  The 


28 


MORPHOLOGY. 


fissure  separating  the  claws  is  the  sulcus  semiannularis  (Figs-  25  and  26).  In  conse- 
quence of  the  subsequent  strong  development  of  the  frontal  and  temporal  lobes,  which 
approach  each  other  more  and  more,  the  angle  formed  by  the  two  limbs  of  the  gyrus  olfac- 

torius  lateralis  becomes  progressively 
more  acute,  although  the  demarcation 
of  the  gyrus  from  the  insula  is  still 
distinct.  In  the  later  stages,  the  limbs 
become  more  closely  approximated  and 
the  apical  part  of  the  convolution  is 
incised  by  the  sulcus  centra/is  insulae, 
which  meanwhile  has  been  formed.  The 
result  is,  that  the  previous  connection 
of  the  two  limbs,  as  well  as  the  de- 
marcation of  the  convolution  towards 
the  insula,  vanishes,  the  convolution 
now  appearing  to  expand  within  the 
substance  of  the  insula. 

Since  these  relations  persist   in  the 
adult,  it   was    assumed    that    the    lateral 

olfactory  convolution,  which  bounds  the  insula  medially,  belonged  to  the  island  ; 
hence  it  was  designated  the  limen  insulae.  The  latter,  however,  belongs  to  the 
rhinencephalon  and  represents  the  gyrus  olfactorius  lateralis,  which  is  subdivided  into  a 


27.  —  Schematic   representation    of   the   gyrus    olfactorius 
lateralis  in  the  foetal  brain. 


Gyrus         Snlctts  Gyrns 

amblens  semiannularis  semilunaris 

FIG.  28. — Oblique  mesial  aspect  of  the  brain  of  a  human  foetus  of  four  months.     Photograph. 

front  and  hind  limb — pars  anterior  and  posterior.      The  area    enclosed  by   the  limbs    was 
named  by  Retzius,   the  anguhis  gyri  olfactorii  -lateralis. 

The   pars   anterior   usually   appears   as    a   fairly   broad   convolution,    which    extends 
from  the  tuberculum   olfactorium   outward  and  somewhat  obliquely  backward  and  is  sep- 


LOBUS   OLFACTORIUS    POSTERIOR.  29 

arated  from  the  substantia  perforata  anterior  by  a  fissure,  the  sulcus  arcuatus  rhinen- 
cephali,  that  follows  the  gyrus  olfactorius  lateralis  medially  as  far  as  the  gyrus  hippocampi. 

Laterally  and  in  front,  the  pars  anterior  joins  the  orbital  convolution  to  form  the 
gyrus  olfactorio-orbitalis  of  Retzius,  which  medially  is  bounded  by  the  postero-lateral 
branch  of  the  sulcus  olfactorius.  The  gyrus  is  commonly  simple,  but  may  be  divided 
into  iwo  parts  by  a  short  fissure;  likewise,  it  may  be  subdivided  by  a  longitudinal 
fissure  into  two  subsidiary  convolutions,  an  anterior  and  a  posterior. 

The  stria  olfactoria  lateralis  passes  as  a  white  fibre-bundle  outward  along  the  pars 
anterior  toward  the  angulus  gyri  olfactorii  lateralis,  here  lying  quite  near  the  substantia 
perforata  anterior.  It  then  bends  backward  in  the  angle  and  later  disappears.  Occa- 
sionally the  lateral  olfactory  root  consists  of  two  bundles,  of  which  the  medial  follows 
the  border  of  the  substantia  perforata,  until  it  is  finally  lost  within  the  substance.  It  is 
to  be  further  noted,  that  a  third  or  middle  root  may  be  found  between  the  lateral  and 
medial  ones.  It  soon  vanishes,  however,  within  the  substantia  perforata. 


Piss 
hippocampi 


Anterior 
end  of  temporal  lobe 

FIG.  29. — Region  around  the  tip  of  the  temporal  lobe  of  the  adult  brain.     Photograph. 

After  recurving  in  the  angle,  the  lateral  olfactory  convolution,  as  the  hind  limb 
or  pars  posterior,  continues  inward  and  backward  toward  the  front  end  of  the  gyrus 
hippocampi. 

On  examining  more  closely  the  antero-median  surface  of  the  gyrus  hippocampi  in 
the  adult  brain,  one  sees  the  convolutions  in  which  the  posterior  limb  fades  away — 
the  gyrus  semilunaris  medially  and  the  gyrus  ambiens  laterally.  The  gyrus  ambiens 
arches  around  the  gyrus  semilunaris  and  then  is  lost  within  the  uncus. 

B.     LOBUS   OLFACTORIUS   POSTERIOR. 

The  substantia  perforata  anterior  is  an  oblique  quadrangular  field  lying  behind 
the  trigonum  olfactorium,  between  the  latter  and  the  tractus  opticus.  It  exhibits  numer- 
ous small  openings  for  the  passage  of  blood  vessels,  especially  in  its  anterior  part  in  the 
vicinity  of  the  trigonum.  This  front  part  constitutes  the  substantia  perforata  anterior 
proper,  the  gyrus  perforatus  rhinencephali. 


30  MORPHOLOGY. 

The  posterior  part,  bordering  the  tractus  opticus,  differs  from  the  anterior  chiefly 
in  its  lighter  color  and  smoother  surface.  This  part  is  known  as  the  diagonal  band  of 
Broca,  or  the  gyrus  diagonalis  rhinencephali. 

Gyrus  perforatus  and  gyrus  diagonalis  form  the  essential  features  of  the  lobus 
olfactorius  posterior.  To  the  latter  belongs  an  additional  small  field  on  the  medial  sur- 
face of  the  hemisphere,  the  gyrus  subcallosus  of  Zuckerkandl.  This  field  is  easily 
located,  since  it  is  continuous  with  the  diagonal  band  of  Broca,  lying  behind  the  area 
parolfactoria,  separated  from  the  latter  by  the  sulcus  parolfactorius  posterior,  and  in 
front  of  the  commissura  anterior  and  the  lamina  rostralis  (Fig.  18). 

The  gyri  subcallosi  (the  stalks  of  the   callosum  of  Broca)   descend  close  together 

from  the  rostrum  of  the  corpus 
callosum.  They  are  separated  from 
each  other  by  a  medial  furrow, 
the  sulcus  subcallos^^s  medianus  of 
Retzius,  and  form  the  narrow  tri- 

*3r- Dia%fHBlrocTd        gonum  praecommissurale,  which  lies 

in  front  of  the  anterior  commissure 
and  belongs  to  the  lamina  praecom- 
missuralis.  The  latter  is  a  thin 
lamella  that  covers  the  anterior  com- 
missure and  passes  over  into  the 

FIG.  30.— Middle  region  of  the  brain-base  of  a  human   foetus.  34-S        lamina     terminalis.  At     the     lower 

cm.  long.     At    each   side   of    the   chiasma  is   seen  the    substantia   per- 
forata  anterior,  with  the  diagonal  band  of  Broca.     (His.)  border     of     the     precommissural     tri- 

gone,  the  two  gyri  subcallosi  diverge 

at  almost  right  angles  and  proceed,  on  each  side,  outward  and  backward  along  the 
optic  tract  as  a  white  band,  the  diagonal  band  of  Broca,  to  reach  the  front  end  of  the 
gyrus  hippocampi. 

Broca' s  band  is  distinguished  by  its  lighter  color  from  the  deeper  tinted  sub- 
stantia perforata  anterior  ;  further,  the  disposition  and  form  of  the  vascular  foramina  are 
characteristic.  These  are  oval  or  elliptical,  their  longer  diameters  paralleling  the  axis 
of  Broca' s  band.  Although  the  band  is  always  present,  it  is  not  always  plainly  visible, 
in  some  cases  it  being  superficial  only  in  certain  places,  while  In  others  it  is  buried 
beneath  a  layer  of  gray  substance  that  must  be  removed  before  the  band  is  seen. 


2.     GYRUS    FORNICATUS. 

To  the  peripheral  region  of  the  rhinencephalon,  the  lobus  olfactorius,  is  attached 
the  central  district.  Here  the  gyrus  fornicatus  first  claims  closer  attention.  It  is  an 
annular  tract  on  the  medial  surface  of  the  hemisphere,  encircled  by  the  cerebral  mantle 
and  composed  of  two  chief  convolutions,  the  gyrus  cinguli  and  the  gyrus  hippocampi, 
connected  with  each  other  by  means  of  the  isthmus. 

The  gyrus  cinguli  forms  the  arching  convolution,  paralleling  the  convex  upper 
surface  of  the  corpus  callosum,  between  the  sulcus  cinguli  and  the  sulcus  corporis 
callosi.  It  presents  numerous  variations,  in  consequence  of  the  inconstant  relations  of 
the  sulcus  cinguli.  The  latter,  in  fact,  does  not  represent  a  simple  fissure,  but  consists 


GYRUS   FORNICATUS.  31 

of  several  parts,  known  as  the  pars  anterior,  pars  intermedia  and  pars  posterior.  As  a 
result,  numerous  annectant  or  bridging  convolutions  arise,  which  unite  the  gyrus  cinguli 
with  the  neighboring  convolutions  of  the  pallium.  When  the  composite  parts  join  to 
form  a  simple  sulcus,  the  course  already  described  (page  22)  as  typical  is  observed. 
In  its  entire  path,  several  incisions,  some  deep,  branch  toward  the  frontal  lobe,  while 
those  passing  into  the  gyrus  cinguli  are  few  and  mostly  short.  The  surface  of  the  gyrus 
cinguli  exhibits  likewise  some  shallow  furrows.  Owing  to  these  peculiarities,  as  well 
as  to  its  smooth  surface,  the  gyrus  is  more  or  less  clearly  defined  from  the  adjacent 
convolutions.  Accordingly,  the  gyrus  cinguli  takes  the  following  course.  It  begins 
narrow  beneath  the  knee  of  the  corpus  callosum,  as  the  direct  continuation  of  Broca's 
field.  In  its  further  course,  around  the  genu  and  over  the  truncus  corporis  callosi,  the 
convolution  is  broader.  Farther  behind,  at  the  bend  around  the  splenium,  it  again 
distinctly  narrows  and,  where  it  is  deeply  incised  by  the  fissura  parieto-occipitalis,  passes 
over  into  the  isthmus  gyri  fornicati. 

When  the  sulcus  cinguli  does  not  form  a  simple  furrow,  the  convolution  assumes 
an  entirely  different  character.  Doubling  and  splitting  of  the  convolution  may  exist,  as 
well  as  separation  into  two,  three  or  four  parts.  Concerning  the  annectant  convolutions, 


Stilciis  corporis 
callosi 


Fissura  pariet 
occipital  is 

Fissura  calcar 


Broca's  field 

Fissura        Fissura         Isthmus 
Fissura  rkinica     Gyr.  hippoc.    %»£         co^ 

FIG.  31. — Medial  cerebral  surface.     Gyrus  fornicatus  is  shaded. 

it  may  be  noted  that  of  these  one  is  fairly  constant  in  the  fore  part  of  the  gyrus 
cinguli  and  establishes  a  connection  between  the  latter  and  the  gyrus  frontalis  superior. 
A  second  annectant  convolution  is  found  in  the  middle  portion,  the  connection  between 
the  gyrus  and  the  lobulus  paracentralis,  while  a  third  appears  in  the  posterior 
division,  providing  continuity  with  the  praecuneus.  The  last  connection  is  often 
double  in  consequence  of  the  sulcus  subparietalis  existing  as  a  separate  furrow  and 
not  as  the  hind  part  of  the  chief  fissure.  In  such  cases  the  gyrus  cinguli  appears  to 
radiate  into  the  praecuneus. 

The  chief  variations  of  the  gyrus  cinguli  are  found  mostly  in  its  front  part. 
Here  the  convolution  may  be  doubled  by  an  inner  or  outer  parallel  fissure. 
If  an  outer  secondary  fissure  be  present,  the  gyrus  cinguli  proper  appears  markedly 
narrowed  at  the  knee  of  the  callosum  ;  in  such  case,  the  convolution  lying  between  the 
secondary  fissure  and  the  sulcus  cinguli  proper  must  be  reckoned  as  part  of  the  gyrus 
cinguli. 


32  MORPHOLOGY. 

The  delimitation  of  the  gyrus  becomes  more  difficult  when  it  consists  of  several 
pieces.  Then  each  portion  behind  appears  as  a  wedge  beneath  the  part  in  front,  and 
the  entire  convolution  is  markedly  narrowed,  particularly  towards  the  genu  corporis  callosi. 
The  convolution  appears  notched  in  its  upper  part.  For  this  reason  Rolando  compared 
it  to  a  cock's  comb  and  called  it  the  "ridged  convolution";  hence  also  the  designation 
of  the  sulcus  cinguli  as  the  "festooned  fissure"  (Pozzi}. 

In  consequence  of  the  deep  incision  into  the  gyrus  fornicatus  by  the  common  limb 
of  the  parieto-occipital  and  calcarine  fissures  behind  the  splenium,  the  isthmus  is  pro- 
duced ;  this  marks  the  transition  of  the  gyrus  cinguli  into  the  gyrus  hippocampi. 

The  gyrus  hippocampi  proceeds  forward,  becomes  broader  and,  at  the  level  of 
the  substantia  perforata  anterior,  bends  around  the  front  end  of  the  fissura  hippocampi 
to  form  the  iincus.  On  its  outer  side,  the  gyrus  hippocampi  is  bounded  by  the 
common  stem  of  the  parieto-occipital  and  calcarine  fissures,  the  anterior  part  of  the 
collateral  fissure  and  the  fissura  rhinica. 

As  the  gyrus  cinguli,  so  also  the  gyrus  hippocampi  exhibits  connections  with  the 
convolutions  lying  to  its  outer  side.  In  this  relation  the  great  variability  of  the  fissura 
collateralis  comes  into  consideration.  When  the  fissura  rhinica  is  connected  with  the 
fissura  collateralis,  two  annectant  convolutions  are  found,  an  anterior  and  a  posterior. 
The  former,  the  gyrus  rhinencephalo-temporalis  anterior,  joins  the  front  part  of  the 
gyrus  hippocampi  with  the  temporal  pole  and  is  one  of  the  most  constant  bridges.  The 
other,  the  gyrus  rhinencephalo-lingualis ',  connects  the  gyrus  hippocampi  with  the  gyrus 
lingualis.  The  last-named  bridge  is  mostly  superficial,  but  may  present  mani- 
fold variations.  It  may  be  divided  by  a  longitudinal  furrow  into  two  parts,  of  which 
one  or  the  other  is  deeply  placed  and  the  remaining  one  is  superficial.  Quite  rarely, 
the  entire  bridge  may  be  deeply  situated,  in  which  case  the  collateral  fissure  ends  in  the 
calcarine.  In  the  event  of  the  fissura  rhinica  being  separated  from  the  fissura  collater- 
alis, a  third  bridge,  the  gyrus  rhinencephalo-fusiformis,  is  present. 

The  surface  of  the  gyrus  hippocampi,  from  where  the  gyrus  approaches  the 
hind  end  of  the  callosum  forward,  particularly  toward  the  bottom  of  the  fissura 
hippocampi,  exhibits  a  lighter  color.  This  tract  is  known  as  the  substantia 
reticularis  alba  (Arnold).  Moreover,  mention  must  be  made  of  the  peculiar  character 
of  the  surface  of  that  part  of  the  gyrus  which  lies  between  the  fissura  rhinica  and 
the  fissura  hippocampi.  Here  the  surface  is  covered  with  numerous  small  nodules 
or  wartlike  elevations,  designated  as  verrucae  gyri  hippocampi. 

3.      HIPPOCAMPUS. 

The  hippocampus  or  cornu  Ammonis  also  belongs  to  the  central  region  of  the 
rhinencephalon.  Since  this  structure  is  seen  only  after  the  lateral  ventricle  has  been 
opened,  its  further  consideration  will  be  postponed  (page  43). 

4.  GYRUS  DENTATUS. 

When,  in  order  to  determine  the  depth  of  the  hippocampal  fissure,  the  gyrus 
hippocampi  is  pulled  downward,  one  sees  a  gray  notched  or  corrugated  band,  the 
fascia  dentata  (Tarini)  or  the  gyrus  dentatus  of  Huxley.  Farther  inward  and  over  the 


GYRUS    DENTATUS. 


33 


gyrus  dentatus,  is  seen  a  white  band,  which  passes  from  the  uncus  gyri  hippocampi 
backward ;  this  is  the  fimbria  hippocampi.  In  its  further  course  the  fimbria  is  con- 
tinuous with  the  fornix. 


Induseum  griseum 


—    Fornix  tranmers. 
Fasciola  cinerea 

Gyms  fasciolaris 
Fiss.  hippocampi 
Fimbria 

Gyms  dentatus 
Sulcus  fimbriodentatu* 


FIG.  32. — Relations  of  the  dentate  gyrus.     Gyrus  dentatus  is  red;  fimbria  and  fornix  are  yellow. 

The  gyrus  dentatus  is  separated  from  the  gyrus  hippocampi  by  the  fissura  hippo- 
campi, and  from  the  fimbria  by  the  sulcus  fimbrio-dentatus .  Following  the  dentate 
gyrus  farther  backward,  it  is  seen  to  run  at  first  parallel  with  the  fimbria  to  the 
splenium  corporis  callosi.  Here  the  gyrus  leaves  the  fimbria,  loses  its  incisions  and 
knobs,  becomes  smooth  and,  as  the  fasciola  cinerea,  passes  around  the  callosum  to 
spread  out  over  the  latter  as  a  thin  lamella  of  gray  substance,  the  induseum  griseum. 
In  the  middle,  the  induseum  exhibits  the  striae  longitudinales  mediates  or  striae  Lancisii, 
while  on  each  side,  in  the  sulcus  corporis  callosi,  lies  the  stria  longitudinalis  lateralis  or 


Stria  L 
Taenia  tecta 


Fissura  interhemisphaerica 


Sulcus  corporis  callosi 


FIG.  33. — Induseum  and  striae  longitudins 


upper  surface  of  the  corpus  callosum. 


taenia  tecta  (Fig.  33).  Induseum  and  striae  longitudinales  run  forward  around  the 
knee  of  the  callosum  and  in  their  farther  course  pass  into  the  gyrus  subcallosus,  with 
which,  in  turn,  Broca's  band  (page  30)  joins. 

According  to  most  authors,  the  fasciola  cinerea  constitutes  the  direct  continuation 
of  the  gyrus  dentatus.  As  shown  by  Retzius,  however,  the  gyrus  dentatus  is  not  pro- 
longed directly  into  the  fasciola  cinerea  (Fig.  34).  On  examining  the  area  beneath  the 
splenium  where  the  gyrus  dentatus  leaves  the  fimbria,  a  thin  strand  is  seen  next  the 
fascia  dentata,  which  likewise  sinks  into  the  depth  of  the  sulcus  fimbrio-dentatus 
3 


34  MORPHOLOGY. 

between  the  fascia  dentata  and  the  fimbria.  This  small  cylindrical  strand  was  called  by 
Retzius  the  gyrus  fasciolaris.  It  is  separated  from  the  gyrus  dentatus  by  a  small 
furrow,  the  sulcus  dentato-fasciolaris,  and  forms,  by  union  with  the  pointed  end  of  the 
gyrus  dentatus,  the  fasciola  cinerea  of  the  authors.  The  fasciola  extends  as  a  gray  semi- 
cylinder  strand  around  the  splenium  and,  on  the  surface  of  the  callosum,  continues  as 
a  broad  plate,  the  gyrus  epicallosus  (Retzius)  or  induseiim  griseum. 


Splenium  cor  par  is  callosi 


Gyrus  fasciolaris 

Fascia  dentata 
Gyri  Andreae  Retzii 


FIG.  34. — Gyrus  fasciolaris  and  gyri  Andreae  Retzii. 

Retzius  agrees  with  Zuckerkandl,  that  the  striae  longitudinales  mediales  and 
laterales  correspond  to  local  elevations  of  the  induseum.  In  front,  they  pass  into  the 
gyrus  subcallosus  and  also  in  part,  at  least  so  far  as  the  taenia  tecta  is  concerned,  into 
the  substance  lying  lateral  to  this  gyrus.  Retzius  further  notes,  that  a  portion  of  the 
gray  lamella  covering  the  callosum  branches  off  at  the  posterior  limit  of  the  splenium  to 
continue  on  the  lower  surface  of  the  latter  and  there  form  an  induseum  inferius.  Since 
this  structure  often  resembles  a  convolution,  it  has  been  designated  by  Retzius  as  the 
gyrus  subsplenialis. 

Following  the  gyrus  dentatus  forward,  the  gyrus  hippocampi  being  drawn  down- 
ward, one  perceives  that  the  dentate  gyrus  here  likewise  gradually  separates  from  the 
fimbria  and  then,  after  a  bend  at  right  angles — the  angulus  gyri — passes  as  a  smooth 
slightly  convex  band  onto  the  uncus.  This  band  of  Giacomini,  as  it  is  termed,  passes 
over  the  under  surface  of  the  uncus,  from  the  outer  side  onward  and  somewhat  back- 
ward, and  thence  continues  onto  its  upper  surface.  The  band  courses  from  the  inner 
side  forward  and  outward,  as  far  as  a  thin  sheet  of  medullary  substance,  the  velum 
terminate  (Abbey),  adhering  to  the  uncus.  This  entire  course  is  plainly  exposed  after 
removal  of  the  gyrus  hippocampi. 

Retzius  recognizes  two  divisions  of  the  gyrus  dentatus,  a  longitudinal  and  a 
transverse.  The  former,  proceeding  from  the  angulus  gyri  dentati,  extends  backward 
within  the  fissura  hippocampi.  The  transverse  division,  on  the  contrary,  proceeding 
from  the  angulus,  represents  the  front  end  of  the  convolution.  The  transverse  part — 


GYRUS   UNCINATUS. 


35 


the  limbus  Giacomini — is  further  subdivided  into  a  pars  occulta,  which  lies  buried  in  the 
hippocampal  fissure,  and  a  pars  aperta,  which  is  visible  on  the  upper  surface  of  the 
uncus.  In  front,  the  pars  occulta  is  limited  by  a  furrow  that  corresponds  morphologi- 
cally to  the  end  of  the  fissura  hippocampi.  Behind,  the  limitation  is  usually  less  definite, 
at  times  the  limbus  Giacomini  appearing  to  pass  over  into  this  part. 

Ciafo»iim"s  band 


Cyrus  intraliml 
—  Retains  — 


Cyrus  dentatus 


FIG.  35. — Giacomini's  band.     The  under  surface  of  the  uncus  is  exposed  by  removing  part  of  the  gyrus  hippocampi. 

On  the  portion  of  the  under  surface  of  the  uncus  lying  in  front  of  Giacomini's 
band,  one  distinguishes  two,  often  only  one,  or  occasionally  three,  sulci  and 
included  convolutions,  which  pass  from  the  anterior  limiting  fissure.  These  are 
designated  as  the  sulci  and  gyri  digitati  externi.  Small  tip-like  extensions  of  the 
Giacomini  band  radiate  forward  for  a  short  distance  within  the  sulci  digitati ;  conse- 
quently, this  part  of  the  limbus  appears  more  or  less  festooned.  The  anterior  termi-. 
nation  of  Giacomini's  band  is  as  yet  undetermined. 


5.    GYRUS   UNCINATUS,  GYRUS    INTRALIMBICUS  AND   GYRUS 
FASCIOLARIS. 

According  to  most  authors,  the  uncus  gyri  hippocampi  or  the  gyrus  uncinatus,  is 
the  continuation  of  the  gyrus  hippocampi,  bent  around  the  anterior  end  of  the  hippo- 
campal fissure,  which  extends  as  far  as  the  beginning  of  the  fimbria  and  is  divided  into 
an  anterior  and  posterior  part  by  the  Giacomini  band.  According  to  Retzius,  however, 
the  front  division  of  the  uncus  must  differ  morphologically  from  the  posterior.  He 
regards,  therefore,  the  anterior  part  as  belonging  to  the  gyrus  hippocampi  and 
designates  this  part  alone  as  the  gyrus  uncinatus;  the  region  lying  behind  the 
Giacomini  band  constitutes  the  gyrus  intralimbic^ls  of  Retzius.  This  intralimbic 
gyrus  appears  sometimes  as  a  small  slightly  arched  surface,  sometimes  as  one  or 
several  knobs,  and  occasionally  is  sharply  defined  by  a  fissure  from  the  fimbria  and 
the  gyrus  dentatus.  The  convolution  continues  for  a  short  distance  backward  within 
the  sulcus  fimbrio-dentatus.  Farther  behind  in  the  same  sulcus,  a  gray  .strand  again 


MORPHOLOGY. 


appears.  This  gradually  thickens,  lies  attached  to  the  gyrus  dentatus,  or  separated 
from  the  latter  by  the  sulcus  dentato-fasciolaris,  and,  as  the  gyrus  fasciolaris  of 
Retzius,  passes  around  the  splenium  corporis  callosi. 

6.    GYRI    ANDREAE    RETZII. 

These,  also  known  as  the  callosal  convolutions,  represent  rudimentary  gyri,  which 
appear  as  round  or  oval  elevations  on  the  medial  surface  of  the  gyrus  hippocampi, 
beneath  the  splenium  and  in  the  angle  formed  by  the  dentate  and  hippocampal  gyri. 
They  are  not  constant  and  may  be  little  more  than  mere  suggestions,  or,  when 

strongly  developed,  may 
resemble  a  spirally  wound 
cord.  Zuckerkandl  desig- 
nates them  as  callosal 
convolutions,  and  Giacomini 
reckons  them,  in  view  of 
their  structure,  as  belong- 
ing to  the  hippocampus. 
G.  Retzius  named  the 
convolutions  in  honor  of 
their  discoverer,  Anders 
Retzius,  his  father,  the  gyri 
Andreae  Retzii  (Fig.  34). 
Summary.  Taking 
a  general  survey  of  the 
entire  rhinencephalon  (Fig. 
36),  we  distinguish  a  periph- 
eral and  a  central  region. 

The  peripheral  region  includes  a  front  and  a  hind  part,  the  lobus  olfactorius 
anterior  and  the  lobus  olfactorius  posterior.  The  central  region  embraces  a  large 
annular  tract  on  the  medial  surface  of  the  hemisphere,  and  includes  the  gyrus  fornicatus 
and  the  gyrus  dentatus. 

Peripheral  and  central  regions  are  closely  united  with  each  other,  the  lobus 
olfactorius'  anterior  being  connected  with  the  gyrus  fornicatus  and  the  lobus 
olfactorius  posterior  with  the  gyrus  dentatus.  Moreover,  the  lobus  olfactorius  anterior  is 
connected,  on  the  one  hand,  with  the  gyrus  cinguli  by  means  of  the  gyrus  olfactorius 
medialis  and,  further  along,  the  area  parolfactoria ;  on  the  other  hand,  it  is  joined 
with  the  front  end  of  the  gyrus  hippocampi  by  means  of  the  gyrus  olfactorius 
lateralis.  The  lobus  olfactorius  posterior  is  connected  with  the  gyrus  dentatus  by 
means  of  Broca's  diagonal  band,  the  gyrus  subcallosus  and  the  induseum  covering 
the  corpus  callosum.  As  will  appear  later,  the  olfactory  centre  is  supposed  to  lie 
chiefly  in  the  cortex  of  the  gyrus  hippocampi.  Therefore,  the  impulses  transmitted 
from  the  nasal  mucous  membrane  by  the  fila  olfactoria  must  be  carried  from  the 
bulbus  olfactorius  and  transferred  to  the  central  region  of  the  rhinencephalon.  The 
course  of  this  olfactory  tract  will  best  explain  the  connections  of  the  individual  parts 
of  the  rhinencephalon  (page  144). 


FIG.  36. — Schematic  representation  of  the  regions  of  the  rhinencephalon: 
Yellow — lobus  olfactorius  anterior  and  gyrus  fornicatus;  red — lobus  olfactorius 
posterior  and  gyrus  dentatus. 


LAMINA  TERMINALIS. 


37 


PARS   OPTICA   HYPOTHALA1Y11. 

This  division  of  the  telencephalon  includes  : 
The  lamina  terminalis, 

The  chiasma  opticum,  with  the  tractus  optici, 
The  tuber  cinereum, 
The  infundibulum, 
The  hypophysis. 

Fornix  Foramen  Monroi 

|  / 


Gyms  ctngnli. 


Cyrus  subcallosus"  •— 


Area  parolfactoria     . 
(Broca's  field) 


Massa  intermedia 


Corpus  mamillare 
FIG.  37. — Median  sagittal  section  through  the  lower  part  of  the  brain. 

The  lamina  terminalis,  or  end-plate,  rises  as  a  thin  sheet  in  front  of  the  chiasma 
opticum  and  continues  in  front  of  the  commissura  anterior  and  the  columnae  fornicis. 
Between  it  and  the  chiasma  lies  the  recessus  opticus.  The  thin  lamella  originally  formed 
the  middle  part  of  the  front  wall  of  the  fore-brain  ;  later  it  is  displaced,  lies  more  deeply, 
and  then  forms  the  anterior  wall  of  the  third  ventricle,  in  whose  roof-plate  it  is  continued. 

The  chiasma  opticum  forms  a  white  quadrangular  plate,  from  the  anterior  cor- 
ners of  which  proceed  the  nervi  optici  and  from  the  posterior  corners  the  tractus  optici. 
The  latter  run  as  flattened  cords  outward  and  backward  along  the  hind  border  of  the 
substantia  perforata  anterior ;  after  passing  around  the  pedunculi  cerebri  and,  farther 
along,  above  and  somewhat  lateral  to  the  uncus  gyri  hippocampi,  they  lead  into  the 
region  of  the  metathalamus. 

The  tuber  cinereum  lies  behind  the  chiasma,  bounded  laterally  by  the  optic 
tracts  and  the  cerebral  peduncles  and  behind  by  the  corpora  mamillaria.  This  gray 
elevation  is  a  thin  lamina  and  assists  in  forming  the  floor  of  the  third  ventricle.  Traced 


3g  MORPHOLOGY. 

forward,  it  passes  into  the  lamina  terminalis,  and  in  this  anterior  position  is  pushed  into 
the  ventricle  by  the  chiasrha.  Below,  the  tuber  cinereum  is  continuous  with  a  hollow 
funnel-shaped  structure,  the  inftindibulum,  whose  cavity  is  known  as  the  recessus  infun- 
dibuli.  To  the  end  of  the  infundibulum  is  attached  the  hypophysis  cerebri  or  pituitary 
body,  a  gray  mushroom-shaped  structure,  about  the  size  of  a  bean,  whose  longest  diam- 
eter is  placed  transversely. 

The  hypophysis,  on  being  sectioned,  is  seen  to  be  composed  of  a  larger  anterior 
and  a  smaller  posterior  lobe.  Genetically,  only  the  posterior  lobe  belongs  to  the  brain, 
as  a  ventral  evagination  from  the  diencephalon.  The  lobus  anterior  originates  as  an 
evagination  from  the  embryonal  oral  recess.  In  consequence  of  constriction  and  isolation, 
the  evagination  later  gives  rise  to  the  hypophysial  vesicle,  which  subsequently  trans- 
forms into  the  gland-like  structure  that,  as  the  anterior  lobe,  becomes  united  with  the 
lobus  posterior. 

Further,  at  particular  points  of  the  tuber  cinereum,  one  often  notes  small  evagina- 
tions.  One,  located  mostly  medial  and  immediately  in  front  of  the  corpora  mamillaria, 
has  been  named  by  Retzius  the  eminentia  saccularis,  while  the  smaller  and  lateral  eleva- 
tions are  the  eminentiae  laterales.  In  Fig.  30  the  eminentia  saccularis  is  plainly  seen. 
It  represents,  perhaps,  a  rudiment  of  the  saccus  vasculosus  strongly  developed  in  the 
bony  and  cartilaginous  fishes. 


INTERNAL  CONFIGURATION   OF  THE  TELENCEPHALON. 

The   examination    of   the    inner   configuration   of   the  end-brain   is   carried  out  most 
advantageously  in  the  following  manner.     A  brain  is  laid  on  its  base  and  the  removal  of 

the  hemispheres  is  begun.  This  is  ac- 
complished by  passing  horizontally,  with 
slow  continuous  stroke,  a  long  brain - 
knife  from  the  convex  lateral  surface  of 
the  hemisphere  as  far  as  the  longitudinal 
cerebral  fissure.  In  this  manner  first 
the  right  and  then  the  left  hemisphere 
are  removed,  from  above  downward,  in 
disk-like  pieces  about  one  centimeter 
thick.  The  last  horizontal  section  should 
fall  about  5  mm.  above  the  dorsal  surface 
of  the  corpus  callosum. 

Each  section  distinctly  exhibits  two 
different  substances — the  .white  substance, 
light  in  color  and  situated  in  the  interior, 
and  the  gray  substance,  which  every- 
where encloses  the  former  and  continues 


FIG.   38. — Horizontal    section    through    the    cerebral    hemi- 
spheres, showing  white  and  gray  substance. 


as    a    band    at    the    periphery    (Fig.    38). 

In  the  first  pieces,  the  white  substance 

is  less  voluminous  than  the  gray.  The  deeper  one  cuts,  however,  the  greater 
the  amount  of  white  substance  revealed,  and  in  the  last  section,  passing  imme- 


CEREBRAL   CORTEX. 


39 


diately  above  the  callosum  (Fig.  39),  in  each  hemisphere  is  seen  a  large  white  medul- 
lary field,  the  centrum  semiovale  (Vieussens),  which  peripherally  is  bounded  by  the  gray 
band  representing  the  cerebral  cortex,  the  substantia  corticalis. 


Striae  longit-ud.  medial, 
*.  Striae  Lancisii 


Stria  longitiid    lateral, 
s.   Taenia  tecta 


Radiatio  cor  par.  callosi 
(Pan  frontalii) 


Radiatio  corpor.  call 
(Pars  parietalii) 


Radiatio  corpor.  callosi 

(Pars  temporal,  and 

occipitalis) 


FIG.  39. — Horizontal  section  at   the  level  of  the  corpus  callosum,  showing  radiation  of  callosal  fibres. 


The  substantia  corticalis  is  not  everywhere  equally  developed  as  to  thickness,  in 
this  respect  varying  according  to  the  region  of  the  brain.  In  general,  the  cerebral 
cortex  is  more  developed  on  the  summit  of  the  convolutions  and  less  so  at  the  bottom 
of  the  fissures,  being  thicker  on  the  outer  convex  surfaces  than  oh  the  medial  and  basal 
aspects  of  the  hemispheres.  The  cortex  reaches  its  maximum 
development  in  the  upper  part  of  the  central  convolutions  and  in 
the  lobus  paracentralis,  and  its  minimum  in  the  occipital  pole. 
When  closely  examined,  even  with  the  unaided  eye,  one  recognizes 
that  the  cerebral  cortex  is  not  homogeneous,  but  is  composed  of 
alternating  gray  and  white  strata  arranged  parallel  with  the  surface. 
The  white  bands  are  known  as  Baillarger 's  stripes.  The  cortex 
of  the  occipital  lobe,  particularly  around  the  calcarine  fissure, 
exhibits  this  stratification  quite  distinctly.  Here  three  layers  are 
found,  an  outer  and  an  inner  gray  stratum  and,  between  them,  a 
thin  light  band,  the  stripe  of  Vicq  d1  Azyr  (Fig.  40),  or,  since 

Gennari  first  described  it,  the  stripe  of  Gennari.  The  explanation  of  this  lamellation 
will  be  given  later  by  the  microscopical  examination  of  the  cerebral  cortex  (page  114). 
In  consequence  of  the  removal  of  the  upper  part  of  the  hemispheres,  as  already 
suggested,  the  corpus  callosum  comes  plainly  in  view.  The  dorsal  surface  of  this  bridge 
lies  before  us,  while  on  each  side  it  is  separated  from  the  overlying  medial  surface  of  the 
hemisphere  by  the  sulcus  corporis  callosi. 


FIG.  40. — Vertical  sec- 
tion through  occipital  lobe. 
The  narrow  light  band  is 
the  stripe  of  Gennari. 


MORPHOLOGY. 


The  corpus  callosum,  or  commissura  cerebri  magna,  forms  a  white  medullary 
mass  that  connects  the  hemispheres.  Strands  of  transversely  coursing  fibres,  the  striae 
transversae,  are  seen  on  the  surface  of  the  truncus  or  body  of  the  corpus  callosum. 
They  penetrate  the  wall  of  the  hemispheres  and  form  the  radiatio  carports  callosi  (Fig. 
39).  The  callosal  radiation  includes  an  anterior,  a  middle  and  a  posterior  part.  The 
anterior  portion,  the  pars  frontalis,  belongs  to  the  callosal  knee  or  genu  and  connects  the 
anterior  parts  of  the  frontal  lobes.  In  consequence  of  the  frontal  lobes  projecting  beyond 

the  genu,  the  connecting  fibres  sweep  in 
curves  far  forward  toward  the  frontal  poles, 
forming  a  sort  of  tongs,  the  forceps  an- 
terior. The  middle  portion,  the  pars 
parietalis,  belongs  to  the  body  of  the 
corpus  callosum  and  unites  the  parietal 
and  temporal  lobes  of  the  two  sides. 
The  posterior  portion  belongs  to  the 
hind  segment  of  the  callosal  body  and 
the  splenium  and,  as  the  pars  temporalis 
and  pars  occipitalis,  connects  the  tem- 
poral and  occipital  lobes.  The  callosal 
fibres  arch  far  backward  toward  the  oc- 
cipital poles  and  form  the  forceps .  poste- 
rior. The  induseum  griseum  covers 
the  upper  surface  of  the  corpus  callosum 
as  a  thin  investment,  that  presents  two 
medial  linear  thickenings,  and,  on  each 
side,  a  lateral  one.  The  medial  longitu- 
dinal stripes,  the  striae  longitudinales  me- 
diales  or  striae  of  Lancisii,  are  separated 
by  a  longitudinal  furrow,  the  raphe  cor- 

poris  callosi.    The  lateral  stripes,  situated  within  the  corresponding  sulcus  corporis  callosi, 
are  the  striae  longitudinales  laterales  or  the  teniae  tectae. 

Now  follows  the  opening  of  the  lateral  ventricles.  Such  parts  of  the  hemispheres 
which  still  overlie  the  corpus  callosum  are  removed  as  far  as  the  level  of  the  dorsal  sur- 
face of  the  callosum.  On  separating  these  parts  with  the  fingers,  in  properly  hardened 
brains,  it  is  possible  to  demonstrate  the  radiatio  corporis  callosi,  especially  the  forceps 
anterior  and  posterior.  A  pointed  knife  is  now  passed  through  the  roof  of  the  lateral 
ventricle,  at  the  side  of  the  body  of  the  callosum  and  from  1-2  cm.  behind  the  genu. 
The  incision  is  lengthened  straight  forward  as  far  as  the  level  of  the  genu  of  the  cal- 
losum and  backward,  in  a  slightly  outwardly  convex  curve,  to  a  point  behind  the 
splenium.  By  gradually  widening  the  opening  medially  and  laterally  the  ventricle  is 
exposed. 

THE  LATERAL   VENTRICLE. 

In  each  lateral  ventricle  we  distinguish  three  outpouchings  or  horns,  the  cornu  ante- 
rius,  the  cornu  posterius  and  the  cornu  inferius,  invading  the  frontal,  occipital  and  temporal 
lob(.s  respectively,  and  the  middle  chief  part  or  body,  the  pars  centralis,  uniting  the  three  horns. 


FIG.  41. — Horizontal  section  at  the  level  of  the  corpus 
callosum.  The  heavy  lines  within  the  centrum  semiovale  indi- 
cate the  incisions  made  in  opening  the  lateral  ventricles. 


LATERAL   VENTRICLE. 


41 


The  front  horn,  the  cornu  anterius,  is  bounded  in  front,  partly  below  and  above 
by  the  fibres  of  the  corpus  callosum.  The  radiation  of  the  callosal  knee  closes  the  ante- 
rior horn  in  front  and  in  addition  forms  a  part  of  the  floor.  The  medial  wall  and,  at 
the  same  time,  the  partition  between  the  two  anterior  cornua  are  contributed  by  the  sep- 
tum pellucidum.  The  latter  consists  of  two  thin  plates,  the  laminae  septi  pellucidi,  between 
which  lies  the  completely  closed  cavum  septi  pellucidi.  A  part  of  the  floor  and  the  lateral 
wall  of  the  anterior  horn  are  formed  by  a  gray  protuberance,  the  corpus  striatum.  The 
thickened  front  part  of  the  latter,  which  projects  into  the  anterior  horn,  is  known  as  the 


Anterior  horn 


Corpus  callosn, 


Corpus  striat. 


Sale,  intermed.  (Stria 
terminal*) 


Inferior  horn  (Enti- 
nentia  collateral) 


FIG.  42. — The  lateral  ventricles,  viewed  from  above. 


head,  or  caput ;  passing  backward,  the  striatum  markedly  narrows  and,  as  a  narrow 
tail-like  band,  the  cauda  corporis  callosi,  continues  through  the  pars  centralis  into  the 
cornu  inferius,  of  which  horn  it  contributes  a  portion  of  the  roof. 

The  pars  centralis  is  a  thin  horizontal  cleft,  roofed  in  by  the  radiation  of  the 
callosum.  On  the  floor,  laterally,  is  the  corpus  striatum  ;  next  follows  the  stria  termi- 
nalis or  stria  cornea.  This  structure  forms  the  floor  of  a  groove,  the  sulcus  interme- 
dius,  situated  between  the  corpus  striatum  and  the  adjoining  thalamus.  The  stripe  is 
called  stria  cornea  on  account  of  its  bluish  coloration,  produced  by  the  underlying  vena 
terminalis.  Medial  to  the  stria  terminalis  comes  a  thin  lamella,  the  lamina  affixa,  that 
covers  the  lateral  part  of  the  thalamus,  to  which  it  is  attached.  Farther  medially,  follow 
the  plexus  chorioidcus  ventriculi  lateralis  and  the  dorsal  surface  of  the  part  of  the  fornix 
which  is  free  and  unattached  to  the  callosum. 

Regarding  the  plexus  chorioideus,  it  must  be  especially  emphasized  that  this  struc- 
ture, composed  of  pial  tissue,  really  only  seemingly  lies  within  the  lateral  ventricle.  As 
in  all  other  parts,  so  also  here  the  ventricle  is  lined  with  ependyma  which  invests  the 
choroid  plexus  with  a  thin  epithelial  layer,  the  lamina  chorioidea  epithelialis ;  the  plexus 


42  MORPHOLOGY. 

lies,  therefore,  extraventricular.  Laterally  the  lamina  chorioidea  begins  at  the  lamina 
affixa ;  medially  it  is  continuous  with  the  epithelium  covering  the  fornix  (Fig.  63). 
On  removing  the  plexus  chorioideus,  the  lamina  chorioidea  epithelialis  is  taken  away 
with  it,  the  epithelial  layer  tearing  through  the  medial  border  of  the  lamina  affixa  and 


Lamina  chorioidea  epfthelialis  ventriculi  tertiz 


Lamina  chorioidta 

epithelialis  ventr.  lateral 


FIG.  43. — Frontal  section  of  the  brain  of  a  human  embryo  of  50  mm.  In  the  middle  are  the  thalami,  with  the  third 
ventricle  (III),  whose  roof  is  formed  by  the  lamina  chorioidea  epithelialis  ventriculi  tertii.  On  each  side,  lateral  to  the 
thalamus,  is  the  hemisphere-vesicle  with  the  lamina  chorioidea  epithelialis  of  the  lateral  ventricle  (II). 

along  the  lateral  border  of  the  fornix.  In  these  locations  delicate  white  stripes,  called 
taeniae,  mark  the  lines  of  separation  ;  hence,  the  taenia  chorioidea  and  the  taema 
fornicis  are  distinguished. 

These  taeniae,  evidently,  as  such  do  not  exist  in  the  normal  and  undamaged 
brain ;  they  are,  therefore,  artefacts  as  are  also  the  taenia  fimbriae,  taenia  thalami  and 
taenia  ventriculi  quarti,  to  be  described  later.  Their  true  relations  are  to  be  under- 
stood only  by  reference  to  embryology.  While  the  original  wall  of  the  embryonic 
brain-tube  for  the  most  part  thickens  during  development  and  becomes  nervous  sub- 
stance, in  certain  places,  namely  in  the  roof  of  the  third  and  of  the  fourth  ventricle  and 
along  a  band-like  area  on  the  medial  wall  of  the  hemisphere,  such  conversion  into 
nervous  tissue  never  occurs,  the  brain-wall  there  being  represented  by  only  a  thin  epi- 
thelial plate,  the  lamina  chorioidea  epithelialis.  Where  the  latter  joins  the  typical  wall, 
the  nervous  substance  is  thinned  out  to  a  slender  wedge. 

The  lamina  chorioidea,  moreover,  at  certain  places  undergoes  a  complicated  invagi- 
nation  toward  the  cavity  of  the  ventricles,  accompanied  by  the  superimposed  pial  tissue,  the 
process  leading  to  the  formation  of  the  plexus  chorioidea.  When  later  the  brain-membranes 
are  removed,  as  when,  for  example,  the  plexus  of  the  lateral  ventricle  is  taken  off,  the 
epithelial  lamina  is  also  removed  and  there  remain  only  delicate  linear  borders,  known 
as  the  taeniae,  that  mark  the  torn  edges  along  those  lines  at  which  the  brain-substance 
ioins  the  epithelial  plate. 


LATERAL   VENTRICLE.  43 

The  plexus  chorioideus  ventriculi  lateralis  passes  forward,  becoming  more  deeply 
placed,  toward  the  anterior  cornu.  Here  is  found  the  Y-shaped  foramen  interventricu- 
lare  Monroi,  which  connects  the  two  lateral  ventricles  with  each  other  and  with  the 
third  ventricle.  Behind,  the  choroid  plexus  continues  outward  and  downward  into  the 
inferior  cornu. 

The  hind  horn,  the  cornu  posterius,  forms  a  narrowing  cleft  of  variable  length, 
with  convex  lateral  and  concave  medial  arched  walls.  The  lateral  superior  wall  is  formed 
by  the  fibre-radiation  of  the  corpus  callosum  ;  the  remaining  boundaries  are  contributed 
by  the  medullary  portion  of  the  occipital  lobe.  On  the  medial  wall,  usually  two  longi- 
tudinal ridges  project  into  the  ventricle.  The  upper  and  less  constant  ridge  is  the 
bulbus  cornu  posterius,  and  is  due  to  the  laterally  arching  callosal  fibres — the  forceps 


Induseum  griseitm 


nix  tramvers. 
Fasciola  cinerea 

Gyrns  fasciolaris 
Fiss.  hippocampi 
Fitnbria 

_.  __ dentatus 

Gyrns  intrnlimb. 

Sitlciis  fimbriodentatus 
Fiss.  hippocampi 


FIG.  44. — Relations  of  the  gyrus  dentatus  (red),  the  fimbria  and  the  fornix  (yellow). 

posterior,  which  here  curve  around  the  deeply  incising  fissura  parieto-occipitalis.  The 
lower  and  constant  ridge  is  the  calcar  avis  and  owes  its  existence  to  the  deep  penetra- 
tion of  the  fissura  calcarina. 

The  lower  horn,  the  cornu  inferius,  curves  downward  and  far  forward  in  the 
temporal  lobe,  to  end  blindly  before  reaching  the  tip.  The  roof  is  formed  laterally  by 
the  callosal  radiation  known  as  the  tapetum,  medially  by  the  cauda  corporis  striati  and 
the  stria  terminalis.  The  floor  exhibits  the  eminentia  collateralis ,  a  longitudinal  ridge 
produced  by  the  deep  invagination  of  the  fissura  collateralis.  Behind,  toward  the  pos- 
terior horn,  the  eminence  continues  into  a  triangular,  slightly  convex  field,  the  trigonum 
collalerale.  The  medial  wall  of  the  inferior  horn  is  occupied  by  a  remarkable  semilunar 
curved  protuberance,  the  hippocampus  or  cornu  Ammonis,  for  whose  production  the  deep 
fissura  hippocampi  is  responsible.  It  begins  behind  the  pars  centralis  or  body  of  the 
lateral  ventricle,  in  advance  of  the  front  end  of  the  calcar  avis,  and  extends  in  a  laterally 
convex  curve  downward  and  forward.  Towards  the  anterior  extremity  of  the  inferior  horn, 
the  hippocampus  broadens  and  then  ends  in  several  claw-like  elevations,  the  digitationes. 
hippocampi,  which  vary  in  development,  in  some  cases  being  merely  suggested,  while  in 
others  they  may  number  four  or  five  or  even  seven.  The  indentations  lying  between 
the  digitations  are  called  the  sulci  interdigitales.  The  marginal  portion  of  the  fornix, 
unattached  to  the  corpus  callosum,  the  dorsal  surface  of  which  has  been  mentioned  in 


44 


MORPHOLOGY. 


relation  with  the  pars  centralis  of  the  lateral  ventricle,  continues  backward  and  outwards; 
it  accompanies  the  hippocampus  medially  into  the  inferior  horn.  The  plexus  chorioideus 
ventriculi  lateralis,  which  is  directly  prolonged  from  the  pars  centralis  into  the  inferior 
horn,  where  it  forms  part  of  the  medial  boundary,  is  especially  well  developed — glomus 
chorioideum — at  the  juncture  of  the  pars  centralis  and  the  inferior  cornu.  If  the  plexus 
be  separated  from  the  fimbria,  a  thin  lamina,  the  taenia  fimbriae,  remains.  At  its  front 
end,  the  wall  of  the  inferior  horn  constitutes  a  delicate  occluding  lamella,  which  is 
clothed  with  the  ependyma  and  termed  the  velum  terminate  of  Aeby.  The  fornix  and 
the  hippocampus  now  merit  closer  examination  (Figs.  44,  45  and  46). 

The  fornix  represents  a  paired  structure,  that  extends  in  a  bold  curve  from  the 
uncus  gyri  hippocampi  as  far  as  the  corpora  mamillaria.  From  the  inferior  horn  of  the 
lateral  ventricle  on  each  side,  the  fimbria,  at  first  narrow,  extends  backward  toward 
the  splenium  and  here  passes  into  the  posterior  limit  of  the  fornix,  the  crus  fornicis, 
which  runs  forward  beneath  the  callosum.  The  two  crura  fornicis  with  the  under  surface 


S-ulcus  fiwbriodentattts 
Cyrus  dentat-us 


Gy 

hippocampi 
(Stibiculnm) 


FIG.  45. — Frontal  section  through  the  inferior  horn  of  the  lateral  ventricle;  free-surface  of  the  gyrus  hippocampi  is  on  the 
right.     Ependyma  is  red;  pia  mater  and  plexus  chorioideus  blue. 

of  the  corpus  callosum  form  an  equilateral  triangle,  whose  apex  is  directed  forward. 
The  two  limbs  of  this  triangle  are  connected  by  strands  of  fibres  running  crosswise  and 
constituting  the  fornix  transversus  or  commissura  hippocampi.  The  entire  triangular 
fibre-plate,  also  designated  as  the  psalterium  or  lyra  Davidis,  is  often  separated  from 
the  under  surface  of  the  callosum  by  a  small  cleft,  the  cavum  psalterii,  sometimes  mis- 
leadingly  called  Verga's  ventricle. 

The  crura  of  the  fornix,  which  curve  around  the  posterior  parts  of  the  thalami  and 
pass  toward  the  under  surface  of  the  callosum,  by  their  union  form  the  corpus  fornicis. 
In  its  posterior  part,  the  body  is  attached  to  the  corpus  callosum,  while  it  extends  as  far 
forward  as  the  vicinity  of  the  foramen  interventriculare.  The  under  surface  of  the 
fornix  exhibits  a  median  groove,  the  sulcus  medianus  fornicis.  In  front,  the  corpus 
fornicis  divides  into  two  anterior  columns,  the  columnae  fornicis •,  which,  as  white  cylindrical 
cords,  sweep  downward  in  forwardly  directed  curves,  in  advance  of  the  thalami  and 
behind  the  commissura  anterior,  and  on  each  side  disappear  in  the  hypothalamic  region. 
They  contribute  the  anterior  boundary  of  the  foramen  interventriculare  and  eventually 
end  in  the  corpora  mamillaria. 


THE   HIPPOCAMPUS. 


45 


The  hippocampus  or  cornu  Ammonis  is,  as  already  mentioned,  produced  by 
the  deeply  invaginating  fissura  hippocampi.  These  relations  are  best  understood  by 
examining  a  frontal  section  passing  immediately  behind  the  uncus  gyri  hippocampi  (Figs. 
45  and  46).  It  will  be  seen,  that  the  entire  cortical  formation  is  pushed  in  toward  the 
ventricle  by  the  penetration  of  the  hippocampal  fissure,  thereby  producing,  in  a  sense, 
an  almost  completely  closed  hollow  cylinder,  in  which  lies  the  gyrus  dentatus  as  a  gray 
cord.  The  upper  end  of  the  scrolled  plate  bends  sharply  outward  and  terminates  as  a 
thin  lamella.  This  invaginated  cortex,  protruding  into  the  ventricle,  is  the  hippo- 
campus. Since  the  latter  at  the  same  time  overlies  the  gyrus  hippocampi,  this  con- 
volution is  also  called  the  subiculum  cornu  Ammonis.  The  white  fibre-layer  covering  the 


Corpus  callosum 


Cortex  gyrt 
hippfcampi 


FIG.  46. — Section  across  the  hippocampal  region;  the  posterior  end  of  the  corpus  callosum  is  viewed  from  in  front 
and  below.  Transition  of  the  alveus  and  the  fimbria  (yellow)  into  the  fornix.  The  course  of  the  gyrus  dentatus  (red)  is 
seen  behind  the  splenium  and,  as  the  induseum,  over  the  corpus  callosum. 

ventricular  surface  of  the  invaginated  cortex  is  the  alveus.      At  the  sharp  outward   bend 
of  the  cortical  plate,  the  alveus  becomes  continuous  with  the  fimbria. 

On  following  the  entire  structure  backward  toward  the  splenium  (best  accomplished 
by  making  several  consecutive  vertical  sections  behind  the  uncus),  it  will  be  seen  that 
the  cortical  formation  of  the  gyrus  hippocampi  passes  over  into  the  cortex  of  the  isthmus 
gyri  fornicati  and,  farther  along,  into  that  of  the  gyrus  cinguli.  The  gyrus  dentatus  separates 
from  the  fimbria  and,  as  the  fasciola  cinerea,  passes  around  the  splenium  to  continue  over 
the  corpus  callosum  as  the  induseum.  Alveus  and  fimbria  are  prolonged  into  the  fornix, 
the  alveus  going  into  the  medial  and  the  fimbria  into  the  lateral  part  of  the  fornix. 


46 


MORPHOLOGY. 


GRAY  MASSES  AND  NUCLEI. 

In  addition  to  the  gray  cortex,  the  substantia  corticalis,  other  definite  gray  masses, 
known  as  the  nuclei  or  ganglia  of  the  end-brain,  are  found  within  the  interior  of  the 
hemispheres.  They  are  the  nucleus  caudatus,  the  nucleus  lentiformis,  the  claustrum 
and  the  nucleus  amygdalae,  and  are  constituent  parts  of  the  stem  of  the  telencephalon. 


FIG.  47. — Frontal  or  vertical  section  of  the  brain,  passing  through  the  knee  of  the  corpus  callosum;   on  each  side  of  the 
latter  is  the  anterior  horn  of  the  lateral  ventricle. 


Capsula  interna 


Nucleus  caudatus 


Nucleus  lenticularis 


FIG.  48. — Model  of   the  corpus  striatum  and  the  thalamus.      Nucleus  caudatus   and  lenticularis  are   yellow;    in  front  and 
below  they  are  continuous,  elsewhere,  separated  by  the  capsula  interna. 


The  nucleus  caudatus  forms  the  part  of  the  corpus  striatum  that  has  been 
mentioned  in  connection  with  the  lateral  ventricle.  The  corpus  striatum  is  divided  by 
a  traversing  fibre-mass  into  two  portions,  a  dorsal  and  medial  one,  the  nucleus  caudatus, 
and  a  lateral  one,  the  nucleus  lentiformis.  The  separating  fibre-mass  is  the  capsula 
interna.  The  thickened  front  end  of  the  corpus  striatum,  that  projects  into  the 
anterior  horn  of  the  lateral  ventricle,  and  the  narrow  band,  that  extends  backward 
through  the  pars  centralis  and  into  the  inferior  horn,  belong  to  the  nucleus  caudatus. 
These  are,  therefore,  more  appropriately  called  respectively  the  caput  and  the  cauda 
nuclei  caudati,  than  the  head  and  tail  of  the  corpus  striatum.  The  lateral  edge  of 


CAUDATE   NUCLEUS. 


47 


the  dorsal  surface  of  the  caudate  nucleus  reaches  the  lateral  margin  of  the  lateral 
ventricle,  its  medial  edge  touches  the  stria  terminalis,  and  its  lateral  surface  lies  against 
the  internal  capsule  (Figs.  49  to  52). 


FIG.  49. — Frontal  section  of  the  brain,  through  the  septum  pellucidum,  which  extends  between  the  body  (Corp,  callos.) 
and  the  rostrum  (C.  c.)  of  the  corpus  callosum  and  forms  the  medial  wall  of  the  anterior  horns  of  the  lateral  ventricles. 
The  corpus  striatum  is  partially  divided  by  the  capsula  interna.  Cl,  claustrum. 


FIG.  50. — Frontal  section  of   the  brain,  through   the   tips   of   the   temporal  lobes.      Cc,  corpus    callosum,  lamina   rostralis; 
Co,  commissura  anterior;    C.  ext.,  capsula  externa;    Cl,  claustrum;   C.  extr..  capsula  extrema. 


48 


MORPHOLOGY. 


The  nucleus  lentiformis,  or  nucleus  lenticularis ,  constitutes  a  wedge-shaped  mass, 
whose  base  is  directed  outward  and  the  apex  inward.  It  lies  lateral  and,  at  the  same 
time,  ventral  to  the  nucleus  caudatus  and  thalamus,  separated  from  the  latter  by  the 
internal  capsule.  In  front  and  ventrally,  the  lenticular  nucleus  is  directly  continuous 
with  the  head  of  the  nucleus  caudatus.  Dorsally,  delicate  gray  stripes  connect  the  two 
nuclei;  hence  the  designation  "corpus  striatum  "  applied  to  the  nuclei  conjointly.  The 
nucleus  lentiformis  bounds  the  internal  capsule  laterally  with  its  downward  and  inward 
sloping  medial  surface.  Its  slightly  convex  lateral  surface  is  vertical  and  borders  the 
capsula  externa,  a  thin  white  medullary  lamella  which  is  limited  externally  by  a 


FIG.  51. — Frontal  section  of  the  brain,  passing  through  the  foramen  interventriculare  Monroi.  Fo,  pillars  of  the 
fornix,  between  which  and  the  corpus  callosum  is  seen  a  part  of  the  septum  pellucidum;  -V.  c.,  nucleus  caudatus;  Co, 
commissura  anterior;  C.  ext.,  capsula  externa;  Cl.,  claustrum;  C.  exlr.,  capsula  extrema. 


narrow  scroll-like  sheet  of  gray  substance  known  as  the  claustrum.  The  ventral 
surface  of  the  lenticular  nucleus  is  horizontal  and,  in  the  middle  part,  continuous 
with  the  cortex  of  the  substantia  perforata  anterior.  Two  thin  medullary  sheets, 
more  or  less  parallel  with  the  lateral  surface,  subdivide  the  lenticular  nucleus  into 
three  segments.  The  outer  one,  the  putamen,  exceeds  the  others  both  in  intensity  of 
color  and  size.  The  inner  segments  are  of  paler  color,  smaller,  and  together  form  the 
globus  pallidus. 

In  the  internal  capsule,  which  extends  between  the  nucleus  caudatus  and  the 
thalamus  on  the  medial  side  and  the  nucleus  lentiformis  on  the  lateral  (Fig.  53),  two 
limbs  are  distinguished,  an  anterior  pars  frontalis  capsulae  internae,  between  the  caudate 


THE   INTERNAL   CAPSULE. 


FIG.  52. — Frontal  section  of   the  brain,  passing   through   the  thalamus    and   the  third  ventricle.     C.  ext.,  capsula  externa- 
Cl.,  claustrum;  C.  exlr.,  capsula  extrema;  //,  tractus  opticus;  m,  corpus  mamillare. 


Corpus  callosn 


Cornu  posterins 


Nncl.  candtitiis 


Nucleus  candatus  (Canda) 


FIG    53. — Horizontal  section  of  the  brain. 


5o  MORPHOLOGY. 

and  lenticular  nuclei,  and  a  posterior  limb,  pars  occipitalis  capsulae  internae,  between  the 
lenticular  nucleus  and  the  thalamus.  The  two  limbs  meet  in  a  laterally  opening  angle 
known  as  the  knee,  the  genu  capsulae  internae. 

The  claustrum  constitutes  a  broad  flattened  nucleus,  a  narrow  plate  of  gray  sub- 
stance, which  ventrally  is  somewhat  thickened  and,  more  medially,  joins  the  substantia 
perforata  anterior.  Its  medial  surface  is  smooth  and  bounds  the  thin  capsula  externa. 
The  lateral  surface  presents  small  projections  and  borders  a  white  medullary  sheet,  the 
capsula  extrema,  between  the  claustrum  and  the  cortex  of  the  island. 

The  nucleus  amygdalae  lies  beneath  the  lenticular  nucleus  in  the  extreme  anterior 
segment  of  the  temporal  lobe.  It  is  continuous  with  the  cortex  of  the  gyrus  hippo- 
campi and  of  the  substantia  perforata  anterior. 

SUMMARY   OF   THE   TELENCEPHALON.  ' 

The  telencephalon,  or  end-brain,  forms  the  most  anterior  and  largest  division  of  the 
encephalon  and  comprises  the  hemisphaerium,  and  the  pars  optica  hypothalami. 

A.  The  hemisphaerium  includes  : 

The  pallium  or  the  cerebral-mantle, 

The  rhinencephalon  or  the  olfactory  brain, 

The  stem  of  the  end-brain. 

The  two  hemispheres,  separated  from  each  other  by  the  fissura  longitudinalis  cer- 
ebri,  are  connected  by  the  lamina  terminalis,  the  corpus  callosum,  the  commissura  anterior 
and  the  fornix  transversus. 

The  pallium  exhibits  the  cerebral  lobes  and  convolutions,  separated  by  the  inter- 
vening clefts  and  furrows.  As  fissures  or  total  furrows  are  designated  those  deeply 
incising  chief  furrows,  which  are  early  developed  and  which,  in  consequence  of  their 
deep  penetration,  push  in  the  wall  of  the  ventricle.  To  these  belong :  the  fissura  cerebri 
lateralis,  the  fissura  parieto-occipitalis,  the  fissura  calcarina,  the  fissura  collateralis  and 
the  fissura  hippocampi.  At  the  bottom  of  the  fissura  cerebri  lateralis  lies  the  fossa  cer- 
ebri lateralis,  which  in  a  measure  corresponds  to  a  ventricular  protrusion  of  the  corpus 
striatum.  The  fissura  parieto-occipitalis  corresponds  to  the  bulbus  cornu  posterioris,  the 
fissura  calcarina  to  the  calcar  avis,  the  fissura  collateralis  to  the  eminentia  collateralis, 
while  the  fissura  hippocampi  is  responsible  for  the  production  of  the  hippocampus  within 
the  inferior  horn. 

As  sulci  or  cortical  furrows  are  designated  the  less  deeply  penetrating  grooves 
which  are  confined  more  to  the  surface  of  the  hemisphere. 

The  chief  divisions  of  the  cerebral  mantle  are :  the  lobus  frontalis,  the  lobus  parie- 
talis,  the  lobus  temporalis,  the  lobus  occipitalis  and  the  insula.  The  latter,  however, 
strictly  regarded,  does  not  belong  to  the  cerebral  mantle,  but  to  the  trunk  of  the 
end-brain. 

The  rhinencephalon  falls  into  the  peripheral  and  cortical  regions. 

The  peripheral  region  comprises  the  lobus  olfactorius,  which  in  turn  is  subdivided 
into  the  lobus  olfactorius  anterior  and  posterior. 


SUMMARY   OF   TELENCEPHALON.  51 

The  lobus  olfactorius  anterior  includes  :  — 

The  bulbus  olfactorius, 
The  tractus  olfactorius, 
The  tuberculum  olfactorium, 
The  area  parolfactoria  of  Broca. 

From  the  tuberculum  olfactorium  the  gyrus  olfactorius  lateralis  extends  laterally 
toward  the  fossa  Sylvii,  here  forms  the  angulus  gyri  olfactorii  lateralis,  then  runs  back- 
ward and  ends  as  the  gyrus  semilunaris  and  gyrus  ambiens  at  the  front  border  of  the 
gyrus  hippocampi.  The  gyrus  olfactorius  medialis  extends  medially  from  the  tuberculum, 
its  continuation  forming  on  the  medial  surface  of  the  hemisphere  the  area  parolfactoria 
of  Broca,  which,  in  turn,  is  prolonged  upward  into  the  gyrus  cinguli. 

The  lobus  olfactorius  posterior  claims  the  substantia  perforata  anterior  and  the  diag- 
onal band  of  Broca,  which  latter  passes  into  the  gyrus  subcallosus,  situated  on  the  medial 
aspect  of  the  hemisphere  behind  the  area  parolfactoria. 

The  cortical  region  has  as  its  chief  components :  — 

The  gyrus  fornicatus,  made  up  of  the  gyrus  cinguli.  and  the  gyrus  hippocampi 
with  the  connecting  isthmus. 

The  hippocampus  or  cornu  Ammonis,  pushed  into  the  inferior  horn  of  the  lateral 
ventricle  by  the  hippocampal  fissure. 

The  gyrus  dentatus. 

The  gyrus  uncinatus,  the  gyrus  intralimbicus,  the  gyrus  fasciolaris  and  the  rudi- 
mentary callosal  convolutions  or  gyri  Andreae  Retzii.  Concerning  the  connections  of 
the  peripheral  and  central  regions  consult  pages  145-149. 

The  stem  of  the  end-brain  has  as  its  most  important  part  the  corpus  striatum, 
which  is  separated  by  the  capsula  intcrna  into  the  medially  situated  nucleus 
caudatus  and  the  laterally  placed  nucleus  lentiformis.  The  latter  is  subdivided  by  the 
medullary  laminae  into  the  putamen  and  the  globus  pallidus.  To  the  stem  of  the 
end-brain  belong,  further,  the  claustrum,  separated  from  the  nucleus  lentiformis  by  the 
capsula  externa,  and  the  nucleus  amygdalae,  located  in  the  extreme  front  part  of  the 
temporal  lobe.  .  All  these  nuclei  are  connected  with  the  cortex  of  the  substantia 
perforata  anterior. 

Within  each  hemisphere,  the  lateral  ventricle  expands  into  its  three  horns,  the 
anterior,  posterior  and  inferior,  and  the  uniting  body  or  pars  centralis.  The  two  lateral 
ventricles  communicate  with  each  other  and  with  the  third'  ventricle  through  the  foramen 
interventriculare  or  foramen  of  Monro. 

B.  The  pars  optica  hypothalami  includes  : — 

The  lamina  terminalis, 

The  chiasma  opticum,   with  the  tractus  optici, 

The  tuber  cinereum, 

The  infundibulum, 

The  hypophysis. 


MORPHOLOGY. 


DIENCEPHALON. 

To  the  diencephalon,  s'ometimes  called  the  inter-brain,  on  account  of  its  position 
between  the  end-  and  the  mid-brain,  belong  : 

The  thalamencephalon  and  the  pars  mamillaris  hypothalami. 

The  diencephalon  surrounds  the  third  ventricle.  The  immediate  roof  of  the  latter 
is  formed  by  the  lamina  chorioidea  epithelialis  and  the  tela  chorioidea  ventriculi  tertii, 
which  lies  above  and  fused  with  the  epithelial  sheet.  As  secondary  coverings,  over  the 
tela,  follow  the  fornix  and  the  corpus  callosum. 

The  dissection  of  the  brain  proceeds  in  the  following  manner,  the  display  of  the 
fornix  being  next  undertaken.  To  this  end,  the  callosum  is  cut  through  transversely, 
from  i  to  2  cm.  in  advance  of  the  posterior  border  of  the  splenium.  This  is  best 
accomplished  by  passing  the  knife,  from  the  side  and  horizontally,  above  the  crus 
fornicis  and  then  cutting  through  the  corpus  callosum  from  below  upward  and 
somewhat  obliquely  backward.  The  callosum  is  now  pulled  up,  its  attachment  to 
the  psalterium  severed,  and  separated  from  the  body  of  the  fornix  and,  farther 
forward,  from  the  upper  border  of  the  septum  pellucidum. 


Corpus  callosmn 
Cornu  anterius 


Cavum  septl  pellucidi 
Lamina  septi  pellncidi 
Nncl.  catidatus 
Stria  terminate 

Corpus  fornicis 
Lamina  affixa 
Hippocampus 
Cms  fornicis 
Plexus  chorioid. 
Eminentia  collaterals 
Calcar  avis 
Bulbns  cornu  post. 
Cerebellum 


FIG.  54-— L 


llosum. 


After  removal  of  the  corpus  callosum  it  is  to  be  noted,  how  on  each  side  the  fim- 
bria  ascends  from  the  inferior  horn  and  passes  into  the  crus  fornicis,  how  the  crura 
fornicis  approach  each  other  and  meet  to  form  the  corpus  fornicis,  and  how  the  columnae 
fornicis  bend  downward  in  front  of  the  foramen  interventriculare  (Fig.  54).  Further  to 
be  observed  are  the  partition  separating  the  lateral  ventricles,  the  septum  pellucidum, 


TELA   CHORIOIDEA. 


53 


with  the  cavum  septi  between  its  laminae,  and  the  course  of  the  plexus  chorioideus 
ventriculi  lateralis  from  the  inferior  horn  through  the  pars  centralis  as  far  as  the 
foramen  interventriculare.  By  means  of  a  probe  or  bristle  may  be  readily 
demonstrated  the  manner  in  which  the  two  lateral  ventricles  are  connected  by  the 
foramen  of  Monro.  The  latter  marks  the  position  at  which  the  plexus  chorioideus 
ventriculi  lateralis  is  continuous  with  the  choroid  plexus  of  the  third  ventricle.  It 
must  not  be  forgotten,  however,  that  the  plexus  really  lies  extraventricular.  On 
removing  the  choroid  plexus,  the  taenia  chorioidea  and  the  taenia  fornicis  are 
recognizable,  and,  likewise,  the  anterior  part  of  the  thalamus. 


Tela  chorioidea. 
ventriculi  tertil 


Vena  septi  pellncidi 

Vena  terminalis 
Stria  terminalis 

Vena  chorioidea 
Vena  cerebri  int. 

Vena  cerebri  magna 
(Galeni) 


FIG.  55. — Lateral  ventricles,  after  removal  of   the  fornix.     Tela  chorioidea  ventriculi  tertii  is  exposed. 


Beneath  the  fornix  lies  the  tela  chorioidea  ventriculi  tertii  (Fig.  55).  In  order  to 
exhibit  the  latter,  we  proceed  in  the  following  manner :  one  peduncle  of  the  fornix  is 
lifted  and  sectioned  with  a  sharp  knife  medialward  and,  at  the  same  time,  obliquely 
backward,  the  section  being  continued  through  the  hind  end  of  the  corpus  callosum, 
thus  cutting  across  the  pars  occipito-temporalis  of  the  radiatio  corporis  callosi.  A 
similar  section  is  executed  on  the  other  side.  The  posterior  end  of  the  callosum  is  now 
raised  and  turned  forward,  with  the  fornix.  The  latter  is  cut  off  at  the  posterior  margin 
of  the  septum  pellucidum,  where  the  corpus  fornicis  passes  into  columnae  fornicis.  After 
removal  of  the  fornix,  the  tela  chorioidea  lies  free,  beneath  which  the  lamina  chorioidea 
epithelialis  alone  remains  as  the  roof  of  the  third  ventricle.  The  removal  of  the  tela 
chorioidea  is  carried  out  from  in  front  ;  it  is  raised  behind  the  columnae  fornicis  and 
carefully  reflected  backward.  Compare  Figs.  62  and  63  for  orientation.  We  now  pass 
to  the  consideration  of  the  thalamencephalon. 


54 


MORPHOLOGY. 


THALAMENCEPHALON. 

The  thalamus  opticus  (Figs.  56  and  57)  presents  an  ovoid  mass  of  gray  sub- 
stance, with  the  thicker  end  behind.  Its  dorsal  and  medial  surfaces  are  free,  while  its 
lateral  and  ventral  ones  are  fused  with  the  neighboring  structures.  The  dorsal  surface 


Ventriculus  tert. 
Trigonum  habenulae 


FIG.  56. — Lateral  and  third  ventricles  exposed;    the  tela  chorioidea  has  been  removed. 

is  slightly  convex  and  covered  by  a  thin  layer  of  white  fibres,  the  stratum  zonale.  The 
outer  limit  is  formed  by  the  stria  terminalis,  lodged  within  the  sulcus  intermedius ;  the 
medial  boundary  is  a  white  stripe,  the  stria  medullaris ,  •  which  indicates  the  boundary 


Tuberculmn  ant    thalanr 

Lamina  affixa 
Ventricnlns  tert. 

Pulviuar 

Corf  its  ge'nicidat.  laterals 
Corpus  genicidat.  mediate 

Corpora  quadrigemina 


Taenia  thalami 

Massa  intermedia 

Stria  medullaris 
Sulcus  chorioideus 
Trigonitm  habenulae 
Habenvla 

Corpus  pineale 
Frenulum 
Nervus  trochlearis 
Velum  medullare  anterius 


FIG.  57' — Thalamus,  epithalamus  and  metathalamus,  viewed  from  above. 

between  the  dorsal  and  medial  thalamic  surfaces.  A  furrow,  the  sulcus  chorioideus,  runs 
from  before  backward  and  outward  and  lodges  the  plexus  chorioideus  of  the  lateral 
ventricle  (Fig.  57).  At  its  front  end,  the  dorsal  surface  exhibits  a  small  round  elevation, 


THE   THALAMUS. 


55 


the  tuberculum  anterius  thalami ;  behind  is  a  similar  projection,  the  pulvinar.  The 
stria  medullaris,  the  medial  boundary,  widens  behind  into  a  triangular  field,  the  trigo- 
num  habenulae.  From  the  latter  proceeds  medially  a  white  fibre-strand,  the  haben- 
ula, which  in  front  joins  with  the  habenula  of  the  opposite  side  to  form  the  commissura 
habenularum,  while  behind  it  passes  into  a  flattened  structure,  the  corpiis  pineale.  Medially 
the  stria  medullaris  is  continuous  with  the  lamina  chorioidea  epithelialis,  over  which 
spreads  out  the  tela  chorioidea.  On  removal  of  the  latter,  the  epithelial  layer  is  sepa- 
rated from  the  stria  medullaris.  There  remains,  however,  along  the  line  of  transition  a 
delicate  border,  the  taenia  thalami,  which  behind  adheres  to  the  dorsal  surface  of  the 
habenula  and  the  pineal  body  and  is  continuous  with  the  taenia  of  the  opposite  side. 

Fornix  Foramen  Monroi 


Commissura  anterior 


Gyms  cingnli  N 


Cyrus  siibcallosns  '  -~  2. 


Area  par olf actor ia. 
(Br oca's  field) 


Massa  intermedia 


nulariim 


Corpus  pineale 


Corpora  quadri- 
getnina 


Corpus  mamillare 
PIG.  58. — Median  sagittal  section  of  the  lower  part  of  the  brain. 

The  medial  surface  of  the  thalamus  is  vertical  and  contributes  the  lateral  wall  of 
the  third  ventricle.  Its  lower  limit  is  indicated  by  the  sulcus  hypothalamicus  or  sulcus 
Monroi,  that  leads  from  the  foramen  interventriculare  to  the  entrance  of  the  aquaeductus 
cerebri.  The  median  surfaces  of  the  two  thalami  are  united,  about  the  middle,  by 
the  massa  intermedia,  often  called  the  middle  commissure. .  The  ventral  surface  of  the 
thalamus  borders  on  the  hypothalamus,  the  lateral  surface  on  the  capsula  interna  (Fig.  58). 

Behind  the  commissura  habenularum  lies  the  corpus  pineale,  so  called  on  account  of 
its  resemblance  to  a  pine-cone.  It  extends  from  an  outpouching  of  the  dorsal  brain-wall, 
the  most  posterior  part  of  the  roof  of  the  third  ventricle,  and  is  a  small  unpaired  body, 
whose  base  is  directed  forward  and  the  apex  backward.  In  its  anterior  part,  at  the  base 
and  between  the  upper  and  lower  lamellae,  lies  the  small  evagination  from  the  third 
ventricle,  termed  the  recessus  pinealis.  The  upper  lamella  is  continuous  on  each  side 


56  MORPHOLOGY. 

with  the  habenula,  the  commissura  habenularum  forming  the  dorsal  wall  of  the  recess. 
The  lower  lamella  is  prolonged  into  the  posterior  commissure  and  the  quadrigeminal  plate. 
Since  the  lamina  chorioidea  epithelialis  is  attached  to  the  dorsal  surface  of  the  pineal 
body,  a  considerable  pocket  is  left  between  this  surface  and  the  lamina  chorioidea  of  the 
third  ventricle;  this  is  the  recessus  suprapinealis.  Sand-like  granules,  the  brain-sand  or 
acervulus,  are  usually  present  within  the  interior  of  the  pineal  body. 

The  posterior  commissura,  commissura  cerebri  posterior,  is  a  bundle  of  trans- 
versely coursing  fibres  which  projects  into  the  ventricle  and  ventrally  bounds  the 
entrance  of  the  recessus  pinealis.  Its  ventral  surface  defines  the  aditus  ad  aquae- 
ductum  cerebri.  The  commissure  is  best  seen  when  the  posterior  wall  of  the  third 
ventricle  is  viewed  from  in  front  (Fig.  59). 


Corpora  qiiadrigemina     


Stria  medullaris 
Taenia  thalnmi 


______     Corpus  pineale 


Commissura  posterior 
III.   Ventriculus 
Massa  intermedia 


FIG.  59- — -Posterior  wall  of  the  third  ventricle,  viewed  from  in  front. 

Turning  to  the  region  behind  the  thalamus,  two  small  protuberances,  the  corpora 
geniculata,  are  to  be  noted  as  additional  parts  belonging  to  the  thalamencephalon.  On 
following  the  tractus  opticus  in  its  course  backward  around  the  cerebral  peduncle,  two 

c Thalamus 

—  Corp.  geniculat.  mediale 

—  Corp.  geniculat.  laterale 
Pednnculus  cerebri    „ 


Tract,  opticus 


PIG.  60. — Course  of  the  tractus  opticus  around  the  cerebral  peduncles  toward  the  corpora  geniculata. 

protuberances  are  encountered — the  elongated  oval  corpus  geniculatum  mediate  and  the 
corpus  geniculafum  laterale.  The  latter  is  a  small  elongated  elevation  at  the  hind  and 
lower  end  of  the  thalamus,  lateral  to  the  pulvinar.  The  medial  body  is  separated  from 
the  lateral  body  and  the  pulvinar  by  a  deep  furrow. 


PARS   MAMILLARIS   HYPOTHALAMI. 

The  pars  mamillaris  hypothalami  comprises  the  corpora  mamillaria.  These,  also 
known  as  the  corpora  catidicantia,  are  two  round  or  oval  relatively  prominent  projections  on 
the  basal  surface  of  the  brain,  between  the  tuber  cinereum  and  the  substantia  perforata 
posterior.  While  separated  from  each  other  by  a  deep  median  cleft,  their  opposed  surfaces 
are  closely  pressed  together  (Fig.  15).  Although  the  boundaries  of  the  mamillary  bodies 


THE   THIRD   VENTRICLE. 


57 


FIG.  6 1. — Part,  of  the  basal  sur- 
face of  the  brain,  showing  the  striae 
albae  tuberis.  (Retzius.) 


are  sharp  medially,  in  front  and  behind,  antero-laterally  each  knob  is  continued  into  a  narrow 
stalk  directed  toward  the  substantia  perforata  anterior.  This  stalk,  the  brachium  corporis 
mamillaris,  is  always  present,  although  variably  developed, 
sometimes  being  broad  and  at  other  times  narrow. 
Occasionally  an  additional  small  lateral  projection,  the 
tubercidum  mamillare  laterale,  is  present,  showing  with 
especial  distinctness  when  it  is  bounded  by  a  small  furrow 
medially  as  well  as  laterally. 

Further,  to  be  mentioned  is  the  stria  alba  tuberis  of 
Lenhosse'k.  This  is  a  delicate  white  band,  scarcely  one 
millimeter  in  width,  that  springs  with  fine  converging  fibres 
at  the  hind  slope  of  the  mamillary  body,  runs  forward,  traverses 
the  tuber  cinereum  obliquely  forward  and  outward,  and, 
finally,  disappears  beneath  the  optic  tract.  According  to 
Lenhosse'k,  the  stria  alba  tuberis  is  nothing  more  than  a 

separated  bundle  of  fornix  fibres,  which  here  pass  superficial  to  the  mamillary  body  (Fig.  61). 
In  several  cases  Retzius  found  the  stria  distinct  only  on  one  side,  while  in  other  cases  it 
was  present  on  both  sides. 

VENTRICULUS   TERTIUS. 

The  third  ventricle  is  a  median  inpaired  cleft-like  cavity,  that  communicates  in  front 
with  the  lateral  ventricles  by  means  of  the  foramen  interventriculare  or  foramen  of  Monra, 
and  behind  with  the  fourth  ventricle  by  means  of  the  aquaeductus  cerebri  or  Sylvian 
aqueduct.  The  front  wall  is  formed,  in  the  lower  part  by  the  lamina  terminalis,  in  the 
upper  part  by  the  commissura  anterior  and  the  columnae  fornicis,  while  the  back  wall 
is  formed  by  the  commissura  habemdarum  and  the  commissura  posterior  (Fig.  58).  The 
side  walls  are  contributed  by  the  medial  surfaces  of  the  thalami  and  of  the  hypothalami, 
separated  by  the  sulci  hypothalamici.  The  floor  of  the  third  ventricle,  in  the  'hind  part, 
is  formed  by  the  cerebral  peduncles  and  the  intervening  posterior  perforated  substance; 
in  the  front  part  it  includes  the  corpora  mamillaria,  the  tuber  cinereum,  with  the  infun- 
dibulum  and  hypophysis,  and  the  chiasma  opticum.  The  immediate  roof  of  the  ventricle 
consists  of  the  lamina  chorioidea  epithelialis,  which  is  fused  with  the  overlying  tela  cho- 

rioidea  ventriculi  tertii,  behind  is  attached  to  the 
dorsal  surface  of  the  habenula  and  of  the  corpus 
pineale,  and  laterally  passes  into  the  stria  medullaris. 
The  tela  chorioidea  ventriculi  tertii,  or 
velum  interpositum,  represents  an  expansion  of 
the  pia  cerebri  between  the  ventral  surface  of  the 
corpus  callosum  and  the  fornix,  on  the  one  hand, 
and  the  dorsal  surface  of  the  diencephalon,  on  the 
other.  The  tela  in  form  resembles  an  equilateral  triangle,  whose  apex  lies  in  front,  behind 
the  columnae  fornicis,  and  whose  base  is  behind,  beneath  the  splenium  corporis  callosi  (Figs. 
55,  62  and  63).  It  consists  of  two  laterally  continuous  sheets,  of  which  the  dorsal  one 
is  attached  to  the  under  surface  of  the  callosum  and  the  fornix,  while  the  ventral  one  in 
the  middle  overlies  the  lamina  chorioidea  epithelialis  of  the  third  ventricle  and  at  the 


FIG.  62. — Tela  chorioidea  ventriculi  tertii  is  blue. 


58  MORPHOLOGY. 

sides  covers  the  larger  part  of  the  dorsal  surface  of  the  thalamus.  Laterally,  where  the 
two  sheets  are  continuous,  richly  vascular  villi-like  tufts  of  the  dorsal  sheet  project  into 
the  lateral  ventricle  to  constitute  the  plexus  chorioideus.  Similarly,  villi  from  the  ven- 
tral sheet  project  into  the  third  ventricle,  where  they  appear  as  two  narrow  stripes  close 
to  the  mid-line  and  together  constitute  the  plexus  chorioidea  ventriculi  tertii.  The 
choroid  plexus  of  the  lateral  ventricle  is  inserted  laterally  in  the  lamina  affixa — taenia 
chorioidea — and  medially  at  the  free  edge  of  the  fornix — taenia  fornicis.  The  two  stripes 
of  the  plexus  chorioidea  ventriculi  tertii  are  attached  laterally  to  the  stria  medullaris — 
taenia  thalami.  The  choroid  plexus  of  the  lateral  ventricles  and  the  band-like  plexus  of 
the  third  ventricle  come  ^together  at  the  foramen  interventriculare.  Between  the  dorsal 
and  ventral  sheets  of  the  tela  chorioidea  lies  arachnoidal  connective  tissue.  In  this  run, 


Corf  11  s  callosnt 


Taenia  f arnica 
Plexus  chorioideus  ventriculi 

lateralis 

Stria  termin.  s.  cornea 

Vena  terminalis 

Lamina  affixa 

Taenia  chorioidea 


Nucleus  caudat. 


Tela  chorioidea  ventriculi 
tertii 


Taenia  thalami 
chorioideus  ventriculi 
tertii 


Ventriculus 
tert. 


FIG.  63. — Diagram  showing   the   relations   of   the   corpus    callosum,  fornix   and   the   tela  chorioidea  ventriculi  tertii.     Pia, 

blue;  ependyma,  red. 

in  the  mid-line  and  close  together,  two  veins,  the  venae  cerebri  internae,  into  which 
empty  in  front  the  vena  septi  pellucidi,  from  the  septum  pellucidum,  the  vena  terminalis, 
from  beneath  the  stria  terminalis,  and  the  vena  chorioidea,  from  the  choroid  plexus  of 
the  lateral  ventricles.  Behind,  at  the  hind  end  of  the  tela  chorioidea,  the  venae  cerebri 
internae  unite  to  form  the  vena  cerebri  magna  of  Galen  (Fig.  55). 

Certain  outpouchings  of  the  third  ventricle  claim  mention.  Of  these  the  recessus 
suprapinealis,  the  recessus  pinealis,  the  aditus  ad  aquaeductum  cerebri,  the  recessus  in- 
fundibuli  and  the  recessus  opticus  have  been  noted.  In  front,  is  the  recessus  triangu- 
laris,  between  the  columnae  fornicis  and  the  commissura  anterior  (Fig.  56). 


THE   NUCLEI    OF   THE   DIENCEPHALON. 

The  Thalamus.  The  thalamus  consists  of  three  chief  nuclei,  the  nucleus  anterior, 
the  nucleits  media/is  and  the  nucleus  lateralis,  which  are  imperfectly  separated  from  one 
another  by  white  medullary  stripes,  the  laminae  medullares. 


NUCLEI   OF   THALAMUS. 


59 


The  nucleus  anterior  includes  the  front  and  dorsal  portion  of  the  thalamus  ;  it 
is,  therefore,  also  known  as  the  dorsal  nucleus.  It  penetrates  wedge-like  between  the' 
medial  and  lateral  nuclei,  is  covered  dorsally  by  the  stratum  zonale,  and  rests  ventrally 
upon  a  bifurcation  of  the  lamina  medullaris  interna.  The  thickened  front  end  produces 
the  protuberance  on  the  dorsal  surface  of  the  thalamus  known  as  the  tuberculum  anterius 
or  corpus  album  subrotundum. 

The  nucleus  medialis  is  bounded  laterally  by  the  lamina  medullaris  interna  and 
medially  by  the  central  gray  substance,  a  sheet  of  gray  matter  which  invests  the  floor 


Caps.   fhaJamus  2  Thafamus 


FIG.  64. — Frontal  section  of  the  brain,  passing  through  the  third  ventricle.  Thalamus  a,  i,  I,  nucleus  anterior,  internus 
and  lateralis  thalami;  Nc,  nucleus  caudatus;  C.  s.,  corpus  subthalamicum;  //,  tractus  opticus;  P.p.,  pes  pedunculi;  m, 
corpus  maniillare;  A,  hippocampus  or  cornu  Ammonis;  C.  exl.,  capsula-externa;  Cl,  claustrum;  C.  extr..  capsula  extrema. 

of  the  third  ventricle  and  the  medial  surface  of  the  hypothalamus  and  also  forms  the 
massa  intermedia  or  middle  commissure.  Anteriorly  the  medial  nucleus  is  closely  con- 
nected with  the  nucleus  anterior,  although  it  does  not  reach  the  front  end  of  the 
thalamus  ;  hence,  in  a  series  of  vertical  sections  carried  through  the  brain,  from  before 
backward,  the  medial  nucleus  first  appears  after  -the  anterior  nucleus  begins  to  diminish. 
Behind,  the  medial  nucleus  passes  into  the  pulvinar. 

The  nucleus  lateralis,  the  largest  of  the  thalamic  nuclei,  includes  the  upper  and 
lateral  portion  of  the  thalamus  and  surrounds,  in  large  part,  the  anterior  and  medial 
nuclei.  Its  medial  boundary  is  formed  by  the  lamina  medullaris  interna ;  laterally  it  is 
bounded  by  the  posterior  limit  of  the  internal  capsule,  from  which  it  is  separated  by  the 


6o 


MORPHOLOGY. 


lamina  medullaris  externa  and  the  stratum  reticulare.  The  dorsal  surface  of  the  nucleus 
is  covered  by  the  stratum  zonale  and  assists  in  forming  the  dorsal  surface  of  the 
thalamus.  The  lateral  part  of  this  last-named  surface  is  clothed  by  the  ependyma  of 
the  lateral  ventricle  and  contributes  that  portion  of  the  floor  of  the  ventricle  known  as 
the  lamina  affixa  ;  the  medial  part  of  the  same  surface  belongs  to  the  external  surface  oi 
the  diencephalon  and  is  covered  by  the  ventral  sheet  of  the  tela  chorioidea.  The 


FIG.  65. — Frontal  section  of  the  brain,  passing  through  the  subthalamic  region.'  Thalamus  a,  i,l,  nucleus  anterior, 
internus  and  lateralis  thalami;  R,  nucleus  ruber;  5.  n.,  substantia  nigra;  P.  p.,pes  pedunculi;  //,  tractus  opticus;  A.cornu 
Ammonis;  Cl,  claustrum;  N.  c.,  nucleus  caudatus. 

ventral  surface  of  the  nucleus  lateralis  rests  upon  the  regio  hypothalamica.  In  front,  the 
lateral  nucleus  aids  the  anterior  one  in  defining  the  foramen  interventriculare ;  behind  it 
passes  into  the  pulvinar. 

The  lamina  medullaris  externa  covers  the  entire  outer  surface  of  the  lateral 
nucleus  and  in  the  region  of  the  pulvinar  broadens  into  a  triangular  medullary  area, 
known  as  Wernicke 's  field  CFig.  207 ). 

The  stratum  reticulare,  or  the  lattice  layer,  forms  the  real  outer  limit  of  the  thalamus 
and  constitutes  a  thin  lamella  of  gray  substance  that  invests  the  entire  outer  surface  of 
the  lateral  nucleus  and  of  the  pulvinar,  separating  the  latter  from  the  internal  capsule. 

As  special  nuclei  of  the  thalamus  are  to  be  noted  the  centrum  medianum  and  the 
nucleus  semilunaris,  the  latter  being  also  known  as  the  corpus  patellare. 

The  centrum  medianum  (Luys)  belongs  to  the  nucleus  medialis  and  presents 
a  rounded  mass  of  gray  substance  that  is  lodged  between  the  medial  and  lateral  nuclei 


SUBTHALAMIC    REGION. 


61 


and  the  pulvinar.  Laterally  it  is  bounded  by  the  lamina  medullaris  interna,  medially 
it  blends  with  the  nucleus  medialis  (Fig.  207). 

The  nucleus  semilunaris  (Flechsig)  belongs  to  the  nucleus  lateralis,  in  whose 
ventral  part  it  lies,  and  leans  against  the  centrum  medianum  in  the  form  of  a  crescent. 

Additional  special  nuclei  of  the  diencephalon  are :  the  nucleus  habenulae  or 
ganglion  habenulae,  within  the  trigonum  habenulae,  and  the  nucleus  corporis  geniculati 
medialis  and  lateralis,  within  the  corresponding  geniculate  bodies. 

Ventral  to  the  thalamus,  the  regie  subthalamica  or  the  hypothalamus  spreads  out 
between  the  internal  capsule  and  the  central  gray  substance  of  the  third  ventricle. 


FIG.  66. — Frontal  section  of  the  brain,  passing  through  the  pulvinar  and  the  upper  end  of  the  Sylvian  aquaeduct.     Co.  p., 
commissura  posterior;  P.  p.,  pes  pedunculi;  A,  cornu  Ammonis  or  hippocampus. 

Within  each  corpus  mamillare  lie  two  nuclei,  a  larger  round  nucleus  medialis 
and  a  smaller  nucleus  lateralis,  which  arches  around  the  medial  nucleus  and  includes 
the  front  and  outer  part  of  the  mammillary  body  (Fig.  201).  Close  to  these  two  nuclei, 
at  the  lateral  and  ventral  side  of  the  nucleus  lateralis,  is  found  a  small  nucleus  accessorius. 

The  nucleus  hypothalamicus,  or  corpus  subthalamicum  (Luys),  lies  within 
the  hind  part  of  the  hypothalamus.  This  lentiform  nucleus  lies  beneath  the  nucleus 
lateralis  thalami  and  medially  to  the  globus  pallidus  of  the  lenticular  nucleus  (Fig.  64). 

The  Capsula  Interna.  Let  us  turn  once  more  to  the  internal  capsule.  It  lies 
between  the  nucleus  lenticularis,  on  the  one  side,  and  the  nucleus  caudatus  and  the 
thalamus  on  the  other.  In  frontal  sections,  it  appears  as  a  lamella  of  white  substance  that 
runs  obliquely  from  above  downward  and  inward,  bounded  externally  by  the  lenticular 


62 


MORPHOLOGY. 


nucleus  and  medially  by  the  caudate  nucleus,  the  thalamus  and  the  subthalamic  region 
(Fig.  67).  An  upper  and  a  lower  region  may  be  distinguished  in  the  internal  capsule. 
The  upper  region,  between  the  lenticular  nucleus  on  the  one  side  and  the  caudate 
nucleus  and  thalamus  on  the  other,  is  known  as  the  regio  thalamica  capsiilae  internae. 
The  lower  region  lies  between  the  nucleus  lenticularis  and  the  hypothalamus  and  is  the 
regio  siibthalamica  capsulae  internae. 


FIG.  67. — Frontal  section  through  the  brain,  showing  the  continuation  of  the  internal  capsule  into  the  pes  pedunculi  (P.p.); 
N.  c.,  nucleus  caudatus;  Cl,  claustrum;   R,  nucleus  ruber;  Py,  pyramidal  tract. 


In  horizontal  sections  (Tig.  68),  the  internal  capsule  forms,  in  the  region  of  thalamus, 
an  outwardly  opening  angle  with  a  shorter  anterior  limb,  pars  frontalis,  lodged  between 
the  lenticular  and  caudate  nuclei,  and  a  longer  posterior  limb,  pars  occipitalis,  between 
the  lenticular  nucleus  and  the  thalamus.  .  The  two  limbs  come  together  at  the  knee,  genu 
capsulae  internae.  The  anterior  limb  of  the  capsule  is  also  called  the  pars  lenticulo-caudata, 
the  posterior  one  the  pars  lenticulo-thalamica.  The  hind  limb  extends  some  millimeters 
beyond  the  nucleus  lenticularis,  this  part  constituting  the  pars  retrolenticularis . 


SUMMARY   OF   DIENCEPHALON. 


The  relations  are  different  in  horizontal  sections  passing"  through  the  subthalamic 
region.  Here,  the  posterior  limb  and  the  pars  retrolenticularis  of  the  internal  capsule 
alone  are  seen,  the  anterior  limb  having  disappeared.  These  relations  are  readily  under- 


Corpits  callosum 


Nucl.  caudatus 


Columna  fornicis 


Ventricnlus  tert. 


Corpus  callc 


Comu  posterius 


FIG.  68. — Horizontal  section  of  the  brain.      The  ir 


Thalamus 


Nucleus  caudatus  (Cauda) 
seen  to  include  two  limbs  and  a  knc 


stood,  when  we  recall  that  in  the  front  part  of  this  region  the  nucleus  lenticularis  is 
continuous  with  the  head  of  the  nucleus  caudatus,  from  which  it  follows,  that  in  the 
subthalamic  region  the  anterior  limb  must  disappear  between  the  lenticular  and  caudate 
nuclei  (Fig.  48 ). 

SUMMARY   OF   THE   DIENCEPHALON. 

The    diencephalon    or   inter-brain    is  subdivided  into   the  thalamencephalon  and  the 
pars  mamillaris  hypothalami. 

A.  The  thalamencephalon  includes : 
The  thalamus, 
The  epithalamus, 
The  metathalmus. 

To  the  epithalaimis  belong  : 
The  corpus  pineale, 

The  regio  habenulae — trigonum  habenulae,   commissura  habenularum. 
The  commissura  posterior. 

To  the  metathalamus  belong : 
The  corpora  geniculata. 


64  MORPHOLOGY. 

B.  The  pars  mamillaris  hypothalami  includes  the  corpora  mamillaria. 
The  thalamus  consists  of  three  chief  nuclei : 
Nucleus  anterior  or  dorsalis, 
Nucleus  medialis    (+   centrum  medianum), 
Nucleus  lateralis  (  +   nucleus  semilunaris). 

The  lateral  boundary  of  the  thalamus  is  formed  by  the  lamina  medullaris  externa  and 
the  stratum  reticulare.  Medially  the  thalamus  is  covered  by  the  central  gray  substance, 
which  likewise  clothes  the  medial  surface  of  the  hypothalamus  and  forms  the  massa 
intermedia. 

Within  the  trigonum  habenulae  lies  the  nucleus  or  ganglion  habenulae. 

The  corpora  geniculata  contain  the  nucleus  corporis  geniculati  medialis  and  lateralis. 

Within  the  hypothalamus,  as  special  centres,  are  found  the  nuclei  of  the  corpora 
mamillaria  and  the  nucleus  hypothalamicus,  or  body  of  Luys. 

The  capsula  interna  lies  between  the  nucleus  lenticularis,  on  the  one  side,  and  the 
nucleus  caudatus  and  the  thalamus,  on  the  other.  It  consists  of  an  anterior  limb,  pars 
frontalis  or  pars  lenticulo-caudata,  .a  posterior  limb,  pars  occipitalis  or  pars  lenticulo- 
thalamica,  with  the  pars  retrolenticularis,  and  the  genu  capsulae  internae.  In  horizontal 
sections  through  the  hypothalamic  region  the  pars  frontalis  is  wanting. 

The  diencephalon  encloses  the  third  ventricle  which  communicates  with  the  lateral 
ventricles  by  means  of  the  foramen  interventriculare,  and  with  the  fourth  ventricle  through 
the  aquaeductus  cerebri. 

The  boundaries  of  the  third  -ventricle  are  as  follows  : — 

Anterior  wall:     Lamina  terminalis, 

Commissura  anterior, 
Columnae  fornicis. 

Posterior  wall:     Commissura  habenularum, 
Corpus  pineale, 
Commissura  cerebri  posterior. 

Lateral  walls :     Medial  surfaces  of  the  thalami  and  the  hypothalami. 

Floor:     Cerebral  peduncles, 

Substantia  perforata  posterior, 

Corpora  mamillaria, 

Tuber  cinereum,  with  infundibulum  and  hypophysis, 

Chiasma  opticum. 

Roof :     Lamina  chorioidea  epithelialis  ; 

secondarily,   tela  chorioidea  ventriculi  tertii, 
fornix  and  corpus  callosum. 

The  diencephalon,  together  with  the  telencephalon,  constitutes  the  prosencephalon 
or  the  fore-brain.  The  pars  optica  hypothalami  and  the  pars  mamillaris  hypothalami 
together  form  the  hypothalamus. 

The  foregoing  relations  are  presented  in  recapitulation  in  the  following  table  :     . 


DERIVATIVES   OF   THE   FORE-BRAIN. 


rf 

?  s 

If 

3  1 

"S   o 

r    ^ 

r~*    5' 

p        i—> 

<   8 

<  2     3 

|  | 

G.'     5* 

1-  1 

1       1 

E    * 

-       P 

g 

1 

§ 

s 

i 

3 

0 

2 

i 

1 

1 

5' 
•o 

3* 

t 

I 

ffi 

VJ 

f 

1 

CO 

DC 

<< 

I 

1 

£• 
3 

M               H 

EJ* 

E- 

^~ 

in 

» 

a? 

I       1 

1 

1 

nr 
5' 

B' 

1-            3 

a 

3 
n 

3 

Regio  habenulae 
Corpus  pineale 
Commissura  post 


Nucleus  anterior 
Nucleus  medialis 
Nucleus  lateralis 


%      3 

<TJ    D- 

I   3" 
1-5 

=  « 


pi 


gd 


Co 
Cl 


!?§:?§: 

5T  g    g    g    g 

1 1 1. 1 
11  s  S 
I  I  8=  S 


66 


MORPHOLOGY. 


MESENCEPHALON. 

The  mesencephalon,  or  mid-brain,  forms  the  smallest  of  the  brain-segments.  Dorsally, 
it  extends  fr»m  the  root  of  the  pineal  body  to  the  posterior  edge  of  the  quadrigeminal 
plate ;  ventrally,  from  the  mammillary  bodies  to  the  front  border  of  the  pons.  It  is  traversed 
longitudinally  by  the  aquae ductus  cerebri  or  Sylvian  aqueduct.  The  dorsal  part  of  the  mid- 
brain  includes  the  quadrigeminal  plate,  lamina  quadrigemina ;  the  ventral  part  the  cerebral 
peduncles,  pedunculi  cerebri  and  the  substantia  perforata  posterior ;  and  the  lateral  part  the 
brachia  quadrigemina. 

LAMINA    QUADRIGEMINA. 

The  quadrigeminal  plate  stretches  from  the  root  of  the  pineal  body  to  the  front 
end  of  the  velum  medullare  anterius.  By  means  of  a  shallow  median  longitudinal 
furrow  and  one  running  transversely,  the  plate  is  subdivided  into  four  parts,  each  of 

which  appears  as  a  white  hemispherical  ele- 
vation. The  two  front  and  larger  elevations 
are  the  anterior  quadrigeminal  bodies,  the 
colliculi  superiores,  the  two  hind  and  smaller 
ones  are  the  posterior  quadrigeminal  bodies, 
the  colliculi  inferiores.  The  front  part  of 
the  longitudinal  furrow,  between  the  superior 
colliculi,  is  broad  and  forms  the  trigonum 
subpineale,  on  which  the  pineal  body  rests  ; 
it  sometimes  presents  a  slight  elevation,  the 
colliculus  subpinealis.  In  its  hind  part,  the 
furrow  is  bounded  by  two  strands  of  white 
fibres,  which  extend  to  the  velum  medullare 
anterius  and  are  known  as  the  frenula  veil 
medullaris  anterioris.  Lateral  from  the  root 
of  the  frenulum,  on  each  side  emerges  the 
trochlear  nerve  (Fig.  69). 

Each  colliculus  continues  laterally  into  an 
arm  or  brachium.  From  the  colliculus  supe- 
rior passes  the  brachium  quadrigeminum  supe- 
rius,  which  runs  as  a  distinct  white  cord  be- 
tween the  thalamus  and  the  medial  geniculate 
body  and  disappears  in  the  vicinity  of  the  lat- 
eral geniculate  body.  The  colliculus  superior, 

brachium  quadrigeminum  superius,  corpus  geniculatum  laterale  and  pulvinar  stand  in  relation 
with  the  tractus  opticus.  From  the  colliculus  inferior  proceeds  the  brachium  quadrigeminum 
inferius,  which  is  broader,  flatter  and  shorter  than  the  upper,  and  disappears  beneath  the 
medial  geniculate  body. 

PEDUNCULI    CEREBRI. 

The  cerebral  peduncles,  with  the  substantia  perforata  posterior,  form  the  ventral 
portion  of  the  mid-brain  and  are  bounded  by  the  optic  tract  in  front  and  by  the  pons 
and  its  peduncles  behind  (Figs.  15  and  75).  Cross-sections  of  the  mid-brain  show  a 


PIG.    69. — The    mesencephalon    and    the    myelencephalon ; 
dorsal  aspect;  IV  ventricle  partially  exposed. 


CEREBRAL   PEDUNCLES. 


67 


subdivision   of    the  cerebral    peduncle  into  a  ventral  segment,   the  basis  pedunculi,   and  a 
dorsal  area,   the  tegmentum.       Between   these  subdivisions  lies  a  grayish   black  substance 


Ventriculus  tert. 
Corpits  geniculat.  med. 

Corpus  genicul.  later  ale  — 
Trigonum  lemnisci  _~ 

Brachium  pontis 
Brachium  conjunctivum 

Crus  cerebelli  ad  pontem 


Stria  mednllar 


FIG.  70. — Dorsal  view  of  mesencephalon  and  myelencephalon.     Schematic. 

Lamina  qitadri gemma 
Aquaeductiis  cerebri 
/  \ 

Sulcits  mesencephali 
lateralis 

Sulcits  mesencephali 

medialis 
s    A7,  ocnlomotorii 

FIG.  71. — Section  through  the  mesencephalon. 
Lamina  quadrigemina 


Sulcits  mesencephali 
lateralis 


Sutstantia  nigra. 


N.  ocnlomotoriiis 


Sulcus  mesencephali  wedialis 
FIG.  72. — Section  through  the  mesencephalon,  at  the  level  of  the  oculomotor  nucleus  (///). 

in  the  form  of  a  crescent,  the  substantia  nigra  of  Sommering.      Superficially  the  basis  and 
the   tegmentum  are   separated   by  two  furrows,   medially  by  the  sulcus  nervi  oculomotcvii 


68  MORPHOLOGY. 

or  sulcus  mesencephali  mediates  and  laterally  by  the  sulcus  mesencephali  latcralis.      Dorsally 
the  tegmentum  is  overlaid  by  the  quadrigeminal   plate. 

The  cerebral  peduncles  emerge  from  the  pons  as  robust  striated  columns  and 
extend  divergingly  toward  the  optic  tracts,  beneath  which  they  disappear.  The  course  of 
the  fibre-bundles  is  worthy  of  note.  They  exhibit  an  outward  and  forward  twist  (Fig. 
75).  Between  the  cerebral  peduncles  lies  the  fossa  interpeduncularis  (Tarini),  whose 
floor  is  formed  by  the  substantia  perforata  posterior,  penetrated  by  numerous  apertures 
for  the  passage  of  blood-vessels.  The  posterior  part  of  the  fossa  deepens  toward  the 
pons  into  the  recessus  posterior,  while  the  anterior  part,  toward  the  corpora  mamillaria, 
sinks  into  the  recessus  anterior.  The  fossa  is  divided  by  a  shallow  median  furrow  into 
two  symmetrical  halves ;  laterally,  toward  the  cerebral  peduncle,  it  is  limited  by  the 
sulcus  nervi  oculomotorii,  from  which  emerge  the  fibre-bundles  of  the  nervus.  oculomotorius. 


Trigonum  lemnisci 


Corpora  q-uadrigemina 

-Pednncuh, 


Tract-us  peduncvlari 


^ 

ANN 

FIG.  73. — Lateral  view  of  the  brain-stem,  showing  tractus  peduncularis  and  taenia  pontis.      Schematic. 

A  special  strand  of  fibres,  the  tractus  peduncularis  transversus,  remains  to  be 
noted.  This  springs  from  the  dorsal  surface  of  the  cerebral  peduncle,  between  the 
brachium  quadrigeminum  posterius  and  the  corpus  geniculatum  mediale,  winds  around 
the  peduncle  midway  between  the  optic  tract  and  the  front  border  of  the  pons  and  dis- 
appears in  the  sulcus  nervi  oculomotorii  (Fig.  75).  According  to  Marburg,  the  tractus 
peduncularis  is  identical  with  the  basal  optic  root  present  in  the  lower  vertebrates.  The 
fibres  are  supposed  to  arise  in  the  retina  and  finally  end  in  the  ganglion  ektomamillare 
located  laterally  to  the  corpus  mamillare. 

AQUAEDUCTUS   CEREBRI. 

This  canal,  the  Sylvian  aqueduct,  forms  a  passage,  lined  with  ependyma,  that  con- 
nects the  third  and  fourth  ventricles.  Dorsally  lies  the  lamina  quadrigemina,  ventrally 
the  tegmentum.  In  cross-sections,  where  the  canal  passes  into  either  ventricle,  it  presents 
an  outline  resembling  a  triangle,  with  the  base  directed  dorsally  and  the  apex  ventrally  ; 
in  the  middle,  its  outline  is  varyingly  cordiform  or  elliptical. 

THE   GRAY   MASSES   OF   THE   MID-BRAIN. 

Surrounding  the  aquaeductus  cerebri  is  the  central  gray  substance,  stratum  gris- 
eum  centrale.  At  the  bottom  of  this  stratum,  at  the  level  of  the  superior  colliculi,  lies 
the  oculomotor  nucleus,  which  joins  the  upward  prolongation  of  the  small  nucleus  nervi 
irochlearis  (Figs.  88  and  89).  Lateral,  at  the  edge  of  the  central  gray  substance,  lies 


SUMMARY  OF  MESENCEPHALON.  69 

the  small  nucleus  radicis  decendentis  nervi  trigemini.  The  nucleus  of  the  posterior 
commissure  and  posterior  longitudinal  bundle  is  located  in  advance  of  that  of  the  oculo- 
motor nerve.  Ventral  and  lateral  to  the  central  gray  substance,  the  formatio  reticularis 
spreads  out.  Between  the  basis  pedunculi  and  the  tegmentum  lies  the  substantia  nigra, 
which  extends  upwards  as  far  as  the  hypothalamus,  while  between  the  substantia  nigra 
and  the  central  gray  substance  is  located  the  red  nucleus,  the  nucleus  ruber  or  nucleus 
tegmenti,  which  appears  round  in  cross-sections  (Fig.  207). 

As  small  nuclei  of  the  tegmentum,  the  ganglion  dor  sale  tegmenti  and  the  ganglion 
profundum.  mesencephali  laterale  et.mediale  are  to  be  noted.  The  ganglion  dorsale  is  a 
small  round  nucleus  lying  behind  the  trochlear  nucleus,  while  the  ganglion  profundum  is 
lodged  within  the  formatio  reticularis,  ventro-lateral  to  the  nuclei  of  the  oculomotor  and 
trochlear  nerves. 

The  anterior  quadrigeminal  body  is  covered  by  the  stratum  zonale  and  contains  the 
stratum  griseum  colliculi  superioris ;  the  posterior  body  encloses  the  centrally  placed 
nucleus  colliculi  inferioris. 

Within  the  posterior  part  of  the  substantia  perforata  posterior,  towards  the  front 
border  of  the  pons,  scattered  nerve-cells  constitute  the  ganglion  interpedunc^dare  of 
Gudden. 

SUMMARY   OF   THE   MESENCEPHALON. 

The  mesencephalon  or  mid-brain  includes  dorsally  the  corpora  quadrigemina,  with 
the  brachia  quadrigemina  and  ventrally  the  pedunculi  cerebri. 

The  superior  colliculi  and  their  brachia,  together  with  the  lateral  corpora  genicu- 
lata,  stand  in  relation  to  the  optic  tracts. 

The  pedunculus  cerebri  is  subdivided  into  the  basis  pedunculi  and  the  tegmentum, 
separated  by  the  substantia  nigra. 

The  chief  gray  masses  are  : — 

The  stratum  griseum  colliculi  superioris, 

The  nucleus  colliculi  inferioris, 

The  stratum  griseum  centrale, 

The  nuclei  of  the  nervus  oculomotorius  and  trochlearis, 

The  small  nucleus  of  the  nervus  trigeminus, 

The  nucleus  of  the  posterior  commissure  and  posterior  longitudinal  bundle, 

The  nucleus  ruber, 

The  substantia  nigra. 

The  smaller  nuclei  are  : — - 

The  ganglion  dorsale  et  profundum  tegmenti, 
The  ganglion  interpedunculare. 

The  mid-brain  is  traversed  by  the  aquaeductus  cerebri  or  Sylvian  aqueduct.  This 
narrow  canal  establishes  communication  between  the  third  and  fourth  ventricles. 

The  mesencephalon  and  the  prosencephalon  together  constitute  the  cerebrum. 


70  MORPHOLOGY. 


ISTHMUS  RHOMBENCEPHALI. 

The  isthmus  rhombencephali  forms  the  transition  from  the  mid-brain  to  rhomb- 
encephalon,  which  latter  is  subdivided  into  the  metencephalon  and  the  myelen- 
cephalon. 

To  the  isthmus  belong  the  brachia  conjunctiva,  the  velum  medullare  anterius  and 
the  trigonum  lemnisci,  which  structures  collectively  constitute  the  dorsal  part  of  the 
isthmus.  Ventral  are  the  cerebral  peduncles.  The  isthmus  surrounds  the  upper  end  of 
the  fourth  ventricle. 

The  brachia  conjunctiva  cerebelli,  crura  cerebelli  ad  cerebrum,  or  superior 
cere  be  liar  peduncles,  form  two  flattened  cylindrical  columns  that  emerge  from  the  cere- 
bellum. They  embrace  the  anterior  medullary  velum,  converge  forward  and  come  together 
behind  the  quadrigeminal  plate.  At  the  sides,  the  brachia  conjunctiva  border  on  the 
pontile  peduncles,  separated  from  the  latter  by  the  sulcus  lateralis  mesencephali,  which 
runs  at  first  toward  the  corpus  geniculatum  lateralis  and  then  laterally. 


, ,  w. r        ^^      Frenuhim  veli  medullar. 

Colliculus  inf. 
Pedunculus  ctrebri  ~^~'^<^^^^^^^/^r~—  Fibrae  arciforme* 

Trigonum  lemnisci  ^\_jfL  jUt^f— \ Velum  medullare  ant. 

Brachium  conjunctiv. 

Krachium  pontis  - 

FIG.  74. — Dorsal  view  of  the  isthmus  rhombencephalon. 

The  velum  medullare  anterius  is  a  thin  medullary  sheet  that  stretches  between 
the  brachia  conjunctiva  or  the  superior  cerebellar  peduncles.  Dorsally  it  is  covered  by  and 
fused  with  the  lingula  of  the  cerebellum  and  assists  in  roofing  in  the  anterior  part  of  the 
fourth  ventricle  (Fig.  79).  From  the  narrow  front  end  of  the  velum  arises  the  frenulum 
veli  medullaris  anterioris  that  extends  toward  the  inferior  colliculi. 

In  advance  of  the  front  end  of  the  brachium  conjunctivum,  lies  a  triangular  field, 
the  trigonum  lemnisci.  It  is  usually  distinguishable  from  the  whiter  brachium  by  its 
gray  color.  Laterally,  the  trigonum  borders  on  the  cerebral  peduncle,  separated  by  the 
sulcus  lateralis  mesencephali,  in  front  it  is  bounded  by  the  brachium  quadrigeminum 
inferius  and  the  inferior  colliculi.  The  area  contains  the  fibre-tracts  of  the  fillet  or  lem- 
niscus  and,  deeply  placed,  the  nucleus  lemnisci  lateralis.  Occasionally  one  notes  delicate 
white  fibre-strands  that  pass  from  the  sulcus  mesencephali  lateralis  over  the  brachium,  par- 
ticularly in  the  vicinity  of  the  quadrigeminal  bodies.  Some  of  the  strands  bend  medially 
at  right  angles  and  pass  backward  through  the  anterior  medullary  velum.  These  fibrae 
arciformes  belong  to  a  bundle  that  ascends  from  the  spinal  cord  to  the  cerebellum,  the 
tractus  spino-cerebellaris  ventralis  or  Gowers'  tract  (page  161). 


PONS   VAROLII. 


METENCEPHALON. 

To  the  metencephalon  belong  the  pons  and  the  cerebellum. 

PONS   VAROLII. 

We  distinguish  a  pars  dorsalis  and  a  pars  basalis  pontis.  The  pars  dorsalis  corresponds 
to  the  pars  intermedia  of  the  floor  of  the  fourth  ventricle.  The  pars  basalis  forms  a  broad 
white  bolster,  that  expands  transversely  and  is  bounded  in  front  by  the  cerebral  peduncles 
and  behind  by  the  medulla  oblongata.  The  lateral  boundary  is  indicated  by  a  line  connect- 
ing the  points  of  emergence  of  the  roots  of  the  trigeminal  and  facial  nerves.  Lateral  to  this 
line,  the  pons  narrows  and  passes  on  each  side  into  the  brachium  pontis,  or  middle  cerebellar 
peduncle,  which  extends  backward  and  enters  the  cerebellum.  The  ventral  surface  of  the 

Tract-us  opticus 
Pedunculus  cerebri 

Tractus  pednncnlaris 
transversus 

Fasc.  superior 
Sulc.  basilaris 


Fasc.  obliquus 

Fasc.  inferior 

Brachium  pontis 

Foramen  caecum 

Fiss.  mediana  ant, 

Sulc.  lateral,  ant 

FIG.  75. — Ventral  view  of  the  brain-stem. 

pons  is  arched  in  the  sagittal  and  transverse  directions  and  exhibits  a  distinct  transverse 
striation.  These  transverse  fibres  are  grouped  in  three  more  or  less  well-defined  bundles. 

The  fasciculus  superior  pontis,  which  courses  in  advance  of  the  attachment  of  the 
trigeminal  nerve. 

The  fasciculus  inferior  pontis,   in  the  lower  third  of  the  pons. 

The  fasciculus  medius  pontis,  between  the  foregoing  bundles,  which  crosses  the 
fasciculus  inferior  in  convex  curves  and  runs  towards  the  places  of  attachment  of  the 
facial  and  acoustic  nerves.  On  account  of  this  course,  the  bundle  is  also  called  the 
fasciculus  obliquus  pontis  or  fasciculus  arcuatus  (Foville). 

The  ventral  pontile  surface  is  modelled  in  the  mid-line  by  a  broad  furrow,  the  sulcus 
basilaris,  in  which  the  basilar  artery  usually  lies.  This  furrow,  however,  is  not  caused  by  the 
basilar  artery,  but  by  the  two  adjacent  longitudinal  ridges,  the  eminentia  pyramidales,  which 
contain  the  pyramidal  tracts.  The  sulcus  basilaris  is  present  even  when  the  basilar  artery 
pursues  an  irregular  course  ;  it  disappears,  however,  in  degeneration  of  the  pyramidal  tracts. 

The  taenia  ponlis  or  fibra  pontis  is  a  special  band  of  fibres  that  arises  in  or  medial  to  the 
sulcus  mesencephali  lateralis,  runs  along  the  front  border  of  the  pons  and  disappears  in 
the  sulcus  ftervi  oculomotorii ;  these  fibres  are  also  called  the  Jila  lateralia  pontis  (Fig.  73). 


72 


MORPHOLOGY. 


THE   CEREBELLUM. 

The  cerebellum,  or  little  brain,  is  a  medially  situated  structure  of  kidney-like  form. 
It  underlies  the  occipital  lobes  of  the  cerebrum,  from  which  it  is  separated  by  the  large 
transverse  fissure,  and  lies  behind  the  pons  and  the  corpora  quadrigemina  and  above  the 
medulla  oblongata.  We  distinguish  an  upper  and  a  lower  surface  and  an  anterior  and  a 
posterior  border.  Both  surfaces  are  arched  ;  the  under  and  more  strongly  convex  surface 
exhibits  in  the  middle  a  broad  sagittal  depression,  the  vallecula  cerebelli,  in  which  lies 
the  medulla  oblongata.  The  anterior  border  is  indented  in  the  mid-line  by  the  incisura 
cerebelli  anterior ;  likewise,  the  posterior  border  by  the  incisura  cerebelli  posterior.  At 
the  borders  of  the  incisura  are  the  anguli  anteriores  and  posteriores.  Front  and  hind 
borders  meet  in  the  anguli  laterales.  The  median  part  of  the  cerebellum,  lying  between 
the  incisura  anterior  and  posterior,  is  known  as  the  worm,  the  vermis  cerebelli.  The 
vermis  superior  is  defined  from  the  lateral  portions,  the  cerebellar  hemispheres,  by  two 
shallow  furrows,  while  the  vermis  inferior  is  more  sharply  demarcated  by  deeper  grooves. 
The  narrow  convolutions,  gyri  cerebelli,  are  separated  from  one  another  by  numerous 
more  or  less  parallel  fissures,  the  sulci  cerebelli,  particularly  in  the  worm  and  the  hemi- 
spheres. A  deeply  penetrating  fissure,  the  sulcus  horizontalis  cerebelli,  extends  on  each 
side  from  the  entrance  of  the  pontile  arm  or  middle  cerebellar  peduncle  in  the  cerebellum 
along  the  front  border  towards  the  angulus  lateralis  and  thence  toward  the  angulus  pos- 
terior. By  means  of  this  fissure,  each  hemisphere  is  divided  into  an  upper  and  a  lower 
surface,  the  fades  superior  and  fades  inferior.  The  sulcus  horizontalis  is  readily  located 
when  we  pass  from  the  position  at  which  the  pontile  arm  enters  the  cerebellum.  The 
sulcus  begins  lateral  to  this  location,  at  first  penetrating  but  slightly,  and  is  here  dis- 
tinguished by  the  narrow  convolutions  of  the  upper  and  lower  surfaces  entering  its 
depth.  From  the  lateral  angle,  the  sulcus  proceeds  as  a  deeper  cleft  along  the  hind 
border,  more  on  the  lower  than  the  upper  surface,  toward  the  incisura  cerebelli  posterior. 

The  worm  and  hemisphere  regions  of  the  cerebellum  are  subdivided  into  definite 
lobes  by  certain  more  or  less  deeply  cutting  fissures.  In  each  hemisphere  three  lobes 
.are  distinguished  :  lobus  superior,  lobus  posterior  and  lobus  inferior,  the  individual  lobes 
of  the  hemisphere  always  corresponding  to  definite  segments  of  the  worm-region. 

A.  Lobus  Superior.  The  lobus  superior  is  bounded  in  front  by  the  incisura 
cerebelli  anterior,  at  the  side  by  the  sulcus  horizontalis  cerebelli  and  behind  by  the  sulcus 

Lingvla  and  Vincula  Ltngulat 


Lobulus 
gnadran- 
gularis 


Pars  ant. 


Pars  post.    - 


Lobulus  centralis  and 
Alae  lobuli  centralis 


Sulc.  superi, 


Lobulus  semilunaris 
superior 


Lobulus  semilunaris 
inferior 


Culmen  \ 

\  Monticnlm 
Declh-e  } 

Sulc.  superior  post 
Folium  vermis 

Tuber  vermis 


FIG.  76. — Upper  surface  of  the  cerebellum. 


THE   CEREBELLUM. 


73 


superior  posterior.  The  sulcus  superior  posterior  starts  in  the  sulcus  horizontals  somewhat 
in  advance  of  the  lateral  angle  and  passes  as  a  deep  curved  fissure,  directed  posteriorly  with 
its  convexity  toward  the  hind  end  of  the  vermis  superior.  The  sulcus  is  readily  recognized 
by  the  different  relations  of  the  bounding  lamellae,  since  those  of  the  superior  lobe  run 
obliquely  outward  and  forward,  while  the  lamellae  of  the  posterior  lobe  run  parallel. 

Passing  from  before  backward,   the  worm  and  the  hemisphere  present  the  following 
parts  of  the  lobus  superior  : 

WORM  HEMISPHERE 

Lingula  ....    Vinculum  lingulae 

Lobulus  centralis Ala  lobuli  centralis 

(  Culmen  )  .    (  Pars  anterior 


Monticulus 


(Decl 


ive 


. .  Lobus  quadrangularis  -j 


Pars  posterior 


The  lingula  lies  deeply  placed  in  the  incisura  cerebelli  anterior  and  consists  ol 
from  four  to  six  or  eight  small  lamellae,  which  rest  upon  and  are  fused  with  the  velum 
medullare  anterius.  Lateral  from  the  posterior  lamellae,  the  vinicula  lingulae  extend 
toward  the  middle  cerebral  peduncle. 

Behjnd  the  lingula,  and  separated  from  it  by  the  sulcus  praecentralis,  follows  the 
lobulus  centralis,  which  overhangs  the  lingula  and  laterally  sends  out  its  lamellae,  the 
aloe  lobuli  centralis. 

The  monticulus,  the  largest  segment  of  the  superior  worm,  lies  behind  the  lobulus 
centralis,  separated  from  the  latter  by  the  sulcus  postcentralis.  It  includes  the  culmen 
and  the  declive  and  corresponds  to  the  hemisphere-segment  of  the  lobtdus  quadrangularis. 
The  latter  is  subdivided  by  the  sulcus  superior  anterior  into  a  pars  anterior  and  a  pars 
posterior,  corresponding  to  the  culmen  and  declive  respectively. 

B.  Lobus  Posterior.  The  lobus  posterior  includes  the  hind  part  of  the  upper 
surface  and  the  posterior  half  of  the  under  surface  of  the  cerebellum.  It  is  separated 


Sulc.  pracpyratnidalls 
Sulc.  postpyramidalis 
Sulc.  inf.  ant. 
Sulc.  inf.  Post.   - 


gracilis 


Lob-uhts  setnilunaris 
inferior 


FIG.  77. — Lower  surface  of  the  cerebellum. 


from  the  lobus  superior  by  the  sulcus  superior  posterior,  and  from  the  lobus  inferior  by 
the  sulcus  postpyramidalis  in  the  worm  and  by  the  sulcus  inferior  anterior  in  the  hemi- 
sphere. The  sulcus  inferior  anterior  may  be  readily  identified  if  the  course  of  the  sulcus 
superior  posterior  be  followed.  It  begins  at  the  side,  on  the  front  border  of  the  hemi- 


74 


MORPHOLOGY. 


sphere,  in  the  sulcus  horizontalis  cerebelli  at  the  place  where  the  sulcus  superior  posterior 
opens,  thence  runs  in  a  curve  toward  the  worm,  where  it  ends  in  the  deeply  pene- 
trating sulcus  postpyramidalis. 

By  means  of  the  sulcus  horizontalis  and  the  sulcus  inferior  posterior,  the  posterior  lobe 
of  the  hemisphere  is  subdivided  into  three  parts,  which  correspond  with  two  segments  of 
the  worm. 

WORM  HEMISPHERE 

Folium  vermis Lobulus  semilunaris  superior 

(  Lobulus  semilunaris  inferior 

Tuber  vermis 3.,    ,    ,  ... 

(  Lobulus  gracihs 

The  folium  vermis  lies  in  the  incisura  cerebelli  posterior,  forms  a  single  stout 
lamella  and  connects  the  two  upper  crescentic  lobules,  the  lobuli  semilunares  superiores. 

The  tuber  vermis  or  tuber  valvulae  corresponds  to  the  lobulus  semilunaris  inferior 
and  the  lobulus  gracilis.  The  lobulus  semilunaris  inferior  is  broad  medially  and  narrow 
laterally,  and  often  separated  into  two  parts,  an  anterior  and  a  posterior,  by  a  lateral 
fissure  that  runs  into  the  sulcus  horizontalis.  The  anterior  and  smaller  part  maintains 
approximately  the  same  width  throughout  and  at  the  side  is  applied  to  the  lateral  end 
of  the  lobulus  gracilis.  The  posterior  and  larger  part  exhibits  usually  two  or  three  small 
lobules,  often  two  crescentic  segments,  of  which  one  begins  medially  at  the  worm  with 
the  thicker  end  and  ends  laterally  in  a  point,  and  the  other  begins  broad  at  the  side 
and  becomes  pointed  toward  the  worm.  The  lobulus  gracilis  lies  in  front  of  the  lobulus 
semilunaris  inferior,  maintains  a  more  or  less  constant  thickness  throughout,  and  is 
separated  from  the  lobulus  semilunaris  inferior  by  the  sulcus  inferior  posterior  and  from 
the  lobus  inferior  by  the  sulcus  inferior  anterior. 

C.    Lobus  Inferior.     This  lobe  includes  the  following  parts  : 

WORM  HEMISPHERE 

Pyramis Lobulus  biventer 

Uvula Tonsilla 

Nodulus Flocculus 


Velum  medullare 
Post. 

Nodulus 


Pyramis 


Lobulus  gracilis 
Lobulus  semilunnris 
inferior 


FIG.  78. — Cerebellum  viewed  from  below  and  in  front. 


The  pyramid,  separated  from  the  tuber  vermis  by  the  sulcus  postpyramidalis \  connects 
the  biventral  lobule  of  the  one  side  with  that  of  the  other.  A  fissure  splits  each  lobulus 
biventer  into  two  portions,  an  anterior  medial  and  a  posterior  lateral. 


Th£   CEREBELLUM. 


75 


Decli 


verm.     Tuber  ver 


Culmen 


Lobulus  central 


The  tonsilla   is  embraced   by  a    medially   concave   curve   described  by  the  sulcus 
praepyramidalis,  which  separates  the  pyramis  from  the  uvula. 

In  advance  of  the  uvula  lies  a  small  conical  structure,  the  nodulus.     Immediately  in 
front  of  the  latter  is  a  thin  white  sheet,  the  velum  medullare  posterius,  that  continues  laterally 
on  each  side  as  the  pedunculi  flocculi  to  join  the  flocculus.     Lateral  to  the  latter,  between 
the     lobulus     quadrangularis     of 
the  superior  lobes  and  the  lobulus 
biventer    is    seen    the    accessory 
flocculus,  flocculus  secundarius. 

On  removing  the  tonsil,  a 
broad  lamella,  the  a/a  uvulae,  or 
the  furrowed  band,  is  seen  pass- 
ing outward  from  the  uvula.  The 
posterior  margin  of  this  band  is 
free,  its  anterior  one  is /continuous 
with  the  posterior  medullary  ve- 
lum. The  deep  recess,  whose  floor 
is  formed  by  the  ala  uvulae  and 
the  velum  medullare  posterius, 
lodges  the  tonsil  and  is  called 

the    nidus   avis.         Its     lateral    Wall        FIG.   79.— Median  sagittal  section   through  the  worm   of   the  cerebellum. 

is    contributed     by    the     lobulus 

biventer  and  the  pedunculus  flocculi,  while  it  is  bounded  medially  by  the  uvula  and 
behind  by  the  pyramid.  The  lobulus  biventer  forms  the  lateral,  the  tonsil  the  medial 
and  the  flocculus  the  anterior  part  of  the  lobus  inferior. 

The  foregoing  relations  are  recapitulated  in  the  following  table: 


IV.  Ventriculus 


Pans 


Medulla  oblongata 


Lobus  superior 


Lobus  posterior 


Lot>us  inferior 


VERMIS 
Lingula  , , 

—  Sulcus  praecentralis 
Lobulus  central  is 

—  Sulcus  postc entrails 


HEMISPHAERIUM 
Vinculum  hngulae 

Ala  lobuli  centralis 


Monticulus 


J  Culmen  ) 
I  Declive  j 


Lobulus   quadran- 
gularis 


Folium  vermis. . . 


Tuber  vermis. 


—  Sulcus  postpyramldalis  — 

Pyramis 

—  Sulcus  praepyramidalis 

Uvula 

Nodulus . . . 


Pars  anterior 

—  Sulc .  sup.  ant.  — 

Pars  posterior 

—  Sulcus  superior  posterior  — 

-         Lobulus  semilunaris  superior 

Sulcus  horizontalis  cerebelli  — 

Lobulus  semilunaris  inferior 

—  Sulcus  inferior  posterior  — 
Lobulus  gracilis 

—  Sulcus  inferior  anterior  — 
Lobulus  biventer 


Tonsilla 

Flocculus  (Flocculus  secundarius) 


On  sectioning  the  cerebellum,  we  recognize  the  internally  situated  white  medullary 
substance,  the  corpus  medullare,  and  the  substantia  corticalis,  which  invests  the  periphery 
as  a  thin  continuous  band  of  gray  matter.  The  medullary  substance  of  the  cerebellum 


76 


MORPHOLOGY. 


is  composed  of  that  of  the  hemispheres  and  of  the  worm,  which  are  continuous  medially. 

Stout    tracts    of    medullary    substance,   the  laminae    medullares,   pass    outward    from    the 

medullary    centre   and   send   off,    mostly   at   acute   angles,    secondary   medullary   laminae. 

The  latter,  in  turn,  give  off  still  smaller  sheets, 
which  finally  are  enclosed  by  gray  substance  and 
represent  the  cerebellar  convolutions  or  folia,  the 
gyri  cerebelli.  This  structure,  when  viewed  in 
sagittal  sections,  is  known  as  the  arbor  medullaris, 
on  account  of  the  tree-like  branching.  In  sagittal 
sections  through  the  worm,  where  this  delicate 
figure  is  particularly  well  seen,  it  is  called  the  arbor 
vitae  vermis. 

The  medulla  of  the  hemispheres  is  con- 
nected with  neighboring  parts  of  the  brain  by 
masses  of  nerve  fibres.  These  masses  constitute 

more  or   less  robust    columns,    which    are    termed   the    peduncles,     crura    or    brachia  of 

the  cerebellum  and   serve   to    connect  it   with  the  pons,  the   mid-brain   and   the    medulla 

oblongata. 

The    brachia    pontis,  middle   cerebellar  peduncles,    or   crura   cerebelli  ad  pontem, 

emerge   on   each   side   from   the    horizontal   sulcus   at   the   anterior   border,    between   the 


FIG.  80.— Schematic  representation  of  the 
crura  cerebelli.  Blue,  superior  peduncle;  green, 
middle  peduncle;  yellow,  inferior  peduncle. 


FIG.  81. — Superior  cerebellar  peduncles,  also  termed   crura   cerebelli   ad  corpora  quadrigemina  and  brachia  conjunctiva. 
Portion  of  the  cerebellum  has  been  removed  to  expose  the  dentate  nuclei. 


lobulus  quadrangularis,    tonsilla   and   flocculus,  and   pass   convergingly  forward,   to   blend 
with  the  pons. 

The  crura    cerebelli   ad  cerebrum  or  superior  cerebellar  peduncles,    also   known 
as  the  crura  cerebelli  ad  corpora  quadrigemina  and  the  brachia  conjunctiva  cerebelli,  lie 


MEDULLA   OBLONGATA. 


77 


in  front  of  the  pontile  crura,  pass  as  flattened  cylindrical  columns  convergingly  forward  and 
disappear  beneath  the  quadrigeminal  bodies.  The  velum  medullare  anterius  stretches  out 
between  them. 


Tractus  cerebro-spinalis 


Brachtutn 
fonti* 


FIG.  82.— Brachia  pontis  or  middle  cerebellar  peduncles  and  corpora  restiforme  or  inferior  cerebellar  peduncles. 

The  crura  cerebelli  ad  medullam  oblongatam  or  inferior  cerebellar  peduncles, 
also  often  called  the  corpora  restiformia,  pass  out  between  the  foregoing  cerebellar  arms 
and  turn  sharply  backward  and  downward  into  the  medulla  oblongata. 


MYELENCEPHALON. 
MEDULLA  OBLONGATA. 

The  upper  boundary  of  the  medulla  oblongata  is  marked  ventrally  by  the 
inferior  edge  of  the  pons  and  dorsally  by  the  striae  acusticae  in  the  floor  of  the 
fourth  ventricle ;  the  lower  boundary  is  indicated  by  the  attachment  of  the  upper 
root-bundles  of  the  first  cervical  nerves,  or,  ventrally,  by  the  lower  limit  of  the 
pyramidal  decussation. 

Let  us  first  examine  the  ventral  surface  of  the  medulla  (Fig.  75).  In  the  mid- 
line  runs  the  fissura  mediana  anterior,  which  is  prolonged  into  the  fissure  of  the  spinal 
cord  bearing  the  same  name,  but  separated  from  it  by  the  crossing  fibre-bundles  tof  the 
pyramidal  decussation,  decussatio  pyramidum.  Toward  the  lower  edge  of  .the  pons,  the 
fissure  widens  into  a  small  depression,  the  foramen  caecum.  On  both  sides  the  median 
fissure  is  bordered  by  the  pyramid,  a  slightly  convex  tapering  column,  broad  above  and 
narrow  toward  the  spinal  cord,  which  appears  to  pass  into  the  anterior  column  of  the 
cord.  Only  a  small  part  of  the  pyramidal  fibres,  however,  actually  maintains  a  course 
along  the  anterior  median  fissure  in  the  anterior  column  of  the  spinal  cord,  since  the 
greater  part  crosses  the  mid-line  in  the  decussatio  pyramidum  and  continues  within  the 
lateral  column  of  the  cord  of  the  opposite  side.  The  part  which  continues  within  the 
anterior  column  is  known  as  the  anterior  pyramidal  tract,  that  within  the  opposite  lateral 
column  as  the  lateral  pyramidal  tract.  These  will  receive  more  detailed  attention  in  the 
consideration  of  the  fibre-paths  (page  161). 


MORPHOLOGY. 


The  pyramid  is  bounded  on  the  outer  side  by  the  sulcus  lateralis  anterior,  from 
which  emerge  the  root-bundles  of  the  hypoglossal  nerve.  Lateral  to  the  sulcus  lateralis 
and  adjoining  the  pyramid  is  seen  the  oliva,  an  ovoid  eminence  whose  thicker  end 
reaches  as  far  as  the  pons  and  which  narrows  below.  The  sulcus  lateralis  anterior  may 
be  marked,  especially  in  its  lower  part,  by  transversely  arching  strands  of  fibres,  known 
as  the  fibrae  arcuatae. 

Turning  now  to  the  dorsal  aspect  of  the  medulla  (Fig.  83),  we  note,  in  the 
lower  part,  the  sulcus  medianus  posterior,  which  above  is  soon  closed  by  a  thin  medullary 
sheet,  the  obex.  At  this  point,  beneath  the  obex,  the  central  canal  of  the  cord  opens 


Fossa  mediana 


Fovea  superior 


Ventricul.  Arantii 


Tuberc.  cinereum 

Funic.  gracilis    Fun.  cuneat.     Fun.  lateralis 
FIG.  83. — Fossa  rhomboidea,  showing  details  of  the  floor  of  the  fourth  ventricle. 


Obex 

Snlc.  median,  post. 
Sulc.  intermed.  post. 
Sulc.  lateral,  post. 


into  the  fourth  ventricle.  Lateral  to  the  sulcus  medianus,  next  comes  the  sulcus  inter- 
medius  posterior,  which  in  the  upper  part  of  the  medulla  runs  laterally  and  then  disappears. 
Farther  outward  is  the  less  distinct  sulcus  lateralis  posterior,  which  likewise  turns  out- 
ward and  may  be  followed  to  about  the  level  of  the  middle  of  the  olive.  Between  the 
median  and  lateral  posterior  sulci,  the  posterior  column,  funiculus  posterior,  represents 
the  upward  prolongation  of  the  corresponding  column  of  the  spinal  cord.  By  means  of 
the  sulcus  intermedius  posterior  the  funiculus  is  subdivided  into  two  special  tracts.  On 
each  side  of  the  posterior  median  fissure,  between  the  latter  and  the  posterior  interme- 
diate fissure,  lies  the  fasciculus  gracilis,  or  Golf  s  column,  continued  upward  from  the 
cord.  In  the  upper  part  it  broadens  into  the  clava  and  then,  again  narrowing,  proceeds 
laterally  and  upward.  Between  the  lateral  and  intermediate  posterior  sulci  runs  the 
fasciculus  cuneatiis,  the  upward  prolongation  of  BurdacK  s  column,  which  at  the  level  of 


THE   FOURTH   VENTRICLE.  79 

the  clava  expands  into  the  tiiberculum  cuneatum  and  higher  up  also  bends  outward. 
Lateral  to  the  sulcus  lateralis  posterior,  between  it  and  the  sulcus  lateralis  anterior,  the 
lateral  column,  funiculus  lateralis,  ascends  from  the  spinal  cord.  After  reaching  the 
lower  end  of  the  olive,  the  column  passes  laterally  and  dorsally,  close  to  the  olive,  almost 
as  far  as  the  pons.  It  is  separated  into  a  dorsal  and  a  ventral  part  by  a  slight  furrow, 
along  which  emerge  the  delicate  root-fibres  of  the  accessory,  vagus  and  glossopharyngeal 
nerves.  The  dorsal  part  of  the  funiculus  lateralis  broadens  above  and,  in  the  region  behind 
the  tuberculum  cuneatum,  swells  into  the  tuberculum  cinereum.  Farther  above,  it  passes 
laterally  in  company  with  the  upper  ends  of  the  column  of  Goll  and  of  Burdach.  These 
upward  and  laterally  directed  portions  of  the  column  of  Goll  and  of  Burdach  and  the 
dorsal  segment  of  the  funiculus  lateralis  collectively  constitute  the  corpus  restiforme  or 
inferior  cerebellar  peduncle,  also  called  the  crus  cerebelli  ad  medullam  oblong  atam,  that 
passes  to  the  cerebellum.  Medially,  the  corpus  restiforme  borders  the  lateral  margin  of 
the  fourth  ventricle.  The  fossa  rhomboidea,  which  forms  the  floor  of  the  fourth  ventricle, 
overlies  the  dorsal  surface  of  the  preceding  parts. 

VENTRICULUS   QUARTUS. 

Isthmus,  metencephalon  and  myelencephalon  together  surround  the  fourth  ventricle, 
a  cavity  filled  with  a  small  amount  of  cerebro-spinal  fluid,  which  below  passes  into  the 
central  canal  of  the  spinal  cord  and  above  is  continuous  with  the  Sylvian  aqueduct. 

Three  segments  are  distinguished,  the  pars  inferior,  the  pars  intermedia  and  the 
pars  superior  ventriculi  quarti. 

The  pars  inferior  belongs  to  the  medulla  oblongata  and  is  embraced  by  the  cor- 
pora restiformia. 

The  pars  intermedia  forms  the  middle  and  broadest  portion  and  continues  above 
into  the  region  between  the  pontile  crura. 

The  pars  superior  belongs  to  the  isthmus  rhombencephali,  its  dorsal  boundary 
being  formed  by  the  brachia  conjunctiva  cerebelli  and  the  velum  medullare  anterius. 

The  floor  of  the  fourth  ventricle  is  formed  by  the  fossa  rhomboidea  and  its  roof 
by  the  anterior  medullary  velum,  the  superior  cerebellar  peduncles  or  brachia  conjunctiva, 
the  posterior  medullary  velum  and  the  tela  chorioidea.  The  posterior  medullary  velum 
and  the  tela  chorioidea  together  constitute  the  tegmen  fossae  rhomboideae,  the  roof  in  the 
limited  sense.  The  edge  along  which  the  anterior  and  posterior  medullary  vela  meet  is 
known  as  the  fastigiiim;  at  this  place  the  fourth  ventricle  projects  into  the  medullary 
substance  of  the  cerebellum,  forming  the  tent-like  recessus  tecti.  The  pars  intermedia 
extends  laterally  on  each  side  into  the  recessus  lateralis  ventriculi  quarti.  Originally  the 
fourth  ventricle  is  a  closed  cavity,  except  above  where  it  communicates  with  the  third 
ventricle  by  means  of  the  aquaeductus  cerebri  and  below  where  it  is  continuous  with  the 
central  canal  of  the  spinal  cord.  Its  floor  and  roof  are  clothed  with  epithelium,  the  epen- 
dyma.  On  the  roof  this  epithelium  lines  the  anterior  and  posterior  medullary  vela  and 
then  continues  as  the  thin  lamina  chorioidea  epithelialis,  which  is  attached  to  the  tela 
chorioidea  ventriculi  quarti  and  thence  is  prolonged  onto  the  borders  of  the  abutting  parts 
of  the  brain.  If  the  ventricle  be  forcibly  opened  behind  from  above,  as  when  the  tela 
chorioidea  is  removed,  the  thin  epithelial  lamina  is  likewise  torn.  The  separation  takes 
place  where  the  lamina  passed  onto  the  more  robust  surrounding  parts  of  the  brain,  only 


8o 


MORPHOLOGY. 


a  thin  white  edge,  the  taenia  ventriculi  quarti,  remaining  along  the  borders  of  the  tear. 
The  taenia  of  the  fourth  ventricle  begins  at  the  obex,  thence  passes  onto  the  corpus  resti- 
forme,  there  forms  the  posterior  border  of  the  recessus  lateralis  and  continues  along 
the  peduncle  of  the  flocculus  and  the  posterior  medullary  velum.  The  tela  chorioidea  of  the 
fourth  ventricle  represents  that  part  of  the  pia  mater  cerebri  that  projects  between  the  ven- 
tral surface  of  the  cerebellum,  more  particularly  the  uvula  and  the  tonsilla,  and  the 
dorsal  surface  of  'the  medulla  oblongata  (Fig.  84).  The  two  pial  sheets  are  united  by 
subarachnoidal  tissue.  The  tela  chorioidea  has  the  form  of  an  equilateral  triangle,  whose 
anteriorly  directed  base  is  attached  in  the  middle  to  the  nodulus  and  at  the  sides  along 
the  posterior  medullary  velum  and  the  flocculus,  and  whose  apex  is  directed  posteriorly 
toward  the  hind  end  of  the  fourth  ventricle.  It  pushes  into  the  ventricle  villiform  proc- 
esses that  constitute  the  plexus  chorioideus  ventriculi  quarti,  subdivided  into  medial  and 


Velum  mednllare  anteriitt 
Corpora  guadrigemina 


IV.  Ventricnlus 


Arachnoidea 
I'elum  medullare  post. 


Tela  chc 


Cisterna  cerebello-medullaris 
Afertura  medialis 


Medulla  oblongata 


FIG.  84. — Sagittal  section  through  fourth  ventricle,  showing  relations  of  the  tela  chorioidez 
Ependyma,  red;    pia  mater,  blue. 


lateral  portions.  The  medial  plexus  consists  of  two  thin  stripes  that  pass  in  the  mid-line, 
close  together,  from  behind  forward  to  the  nodulus.  From  the  latter,  the  lateral  plexus 
continues,  on  each  side,  outward  into  the  recessus  lateralis  ventriculi  quarti.  In  the  early 
condition,  the  tela  chorioidea,  with  the  lamina  chorioidea  epithelialis,  completely  closes 
the  posterior  part  of  the  fourth  ventricle.  Later,  however,  openings  are  formed  at  those 
places,  at  which  the  tela  chorioidea  and  the  lamina  epithelialis  are  broken  through.  Such 
an  opening  is  the  apertura  medialis  ventriculi  quarti  or  the  foramen  of  Magendi,  situated 
in  the  posterior  part  of  the  tela  chorioidea  immediately  in  front  of  the  obex.  At  the 
sides,  in  each  lateral  recess,  is  found  the  apertura  lateralis  ventriculi  quarti  (Key-Retzii) 
or  the  foramen  of  Luschka.  Through  these  three  openings  the  ends  of  the  medial  and 
lateral  parts  of  the  choroid  plexus  of  the  fourth  ventricle  pass  and  project  into  the  sub- 
arachnoid  space,  communication  between  the  ventricle  and  the  subarachnoid  space  being 
in  this  manner  established.  The  villi  which  protrude  through  the  apertura  lateralis  are 
readily  found,  since  they  lie  medial  to  the  flocculus,  between  the  latter,  the  lobulus 
biventer  and  the  tonsilla. 


FLOOR  OF  THE  FOURTH  VENTRICLE. 


81 


Fossa  Rhomboidea. — The  floor  of  the  fourth  ventricle,  the  fossa  rhomboidea,  is, 
as  indicated  by  its  name,  rhomboidal  in  outline.  Its  posterior  part,  bordered  by  the  corpora 
restiformia,  belongs  to  the  myelencephalon;  its  middle  part  lies  in  the  metencephalon;  and 
its  anterior  part  belongs  to  the  isthmus.  By  means  of  a  longitudinal  furrow,  sulcus 
medianus  fossae  rhomboidcae,  it  is  divided  into  symmetrical  halves.  Transversely  coursing 
white  bands,  the  striae  medullares  or  striae  acusticae,  which  run  from  the  lateral 
recesses  toward  the  mid-line,  separate  the  pars  superior  from  the  pars  inferior 
fossae  rhomboideae.  The  part  of  the  fossa  included  between  the  medullary  striae  constitutes 
the  pars  intermedia. 

The  striae  medullares  present  many  variations  in  their  course  and  develop- 
ment. They  may  be  wanting  or  many,  but  are  seldom  identical  in  development 


Fossa  medlana 


Fovca.  superit 


Centrical.  Arantii 


Obex 

Side,  median,  post. 
Sulc.  intermed.  post. 
Futtic.  gracilis     Fan.  cimeat.     Fun.  lateralis  Sulc.  lateral,  post. 

FIG.  85. — Dorsal  surface  of  the  medulla  oblongata.     Fossa  rhomboidea. 

and  course  on  the  two   sides.     Often   they   run    obliquely  outward  and  upward  from  the 
sulcus  medianus. 

The  pars  inferior  of  the  ventricle  deepens  in  its  lower  portion,  presents  several  fields 
denned  by  furrows  and,  on  account  of  its  peculiar  shape,  is  called  the  calamus  scriptorius. 
At  the  lower  border  of  the  pars  inferior  lies  the  obex,  a  thin  white  medullary  sheet  from 
which  the  taeniae  ventriculi  quarti  pass  laterally.  Immediately  in  front  of  the  obex,  where 
the  sulcus  medianus  sinks  into  the  central  canal  of  the  spinal  cord,  is  a  small  depression,  the 
ventriculus  Arantii.  In  the  pars  superior,  the  median  sulcus  widens  into  the  fossa 
mediana.  On  each  side  of  the  median  furrow,  a  flat  ridge,  the  eminentia  mediates,  extends 
the  entire  length  of  the  ventricular  floor.  This  ridge  is  narrow  in  its  lower  part  and  forms 
6 


82  MORPHOLOGY. 

a  triangular  field,  the  trigonum  nervi  hypoglossi,  whose  base  is  above  at  the  striae  medul- 
lares  and  apex  below,  directed  toward  the  ventriculus  Arantii. 

On  careful  inspection,  two  special  divisions  of  this  field  are  recognized,  an  outer 
broader  part,  the  area  plumiformis  (Retzius),  and  an  inner  narrower  one,  the  area 
medialis  trigoni  nervi  hypoglossi  (Retzius).  At  the  border  between  those  two 'fields 
are  to  be  seen  mostly  short,  obliquely  coursing  delicate  furrows  and  folds,  and  likewise 
a  thin  feathery  band.  Such  markings  are  often  visible  also  at  the  lateral  border  of  the 
trigonum  hypoglossi.  Retzius,  therefore,  named  this  lateral  and  broader  field,  ' '  area 
plumiformis. ' ' 

In  the  upper  part  of  the  ventricular  floor,  the  eminentia  medialis  is  broader  and  pro- 
jects more  into  the  ventricle.  The  elevation  is  termed  the  colliculus  facialis.  Laterally,  the 
eminentia  medialis  is  defined  by  the  sulcus  limitans,  which  in  the  pars  superior  widens  into 
the  fovea  superior,  and  in  the  pars  inferior  into  the  fovea  inferior.  Below  the  fovea 
inferior,  and  lateral  to  the  trigonum  hypoglossi,  is  seen  a  gray  oblique  triangular  field, 
known  as  the  dla  cinerca,  which  begins  pointed  at  the  fovea  inferior  and  broadens  toward 
the  lower  border  of  the  fossa  rhomboidea. 

In  front  of  the  posterior  border  of  the  fossa  and  behind  the  ala  cinerea,  lies  a 
small  gray  mammillated  field,  the  area  postrema,  that  extends  from  the  mid-line  along  the 
lower  border  of  the  ventricle  forward  and  outward.  A  light  narrow  band,  known  as  the 
funiculus  separans,  runs  from  the  opening  central  canal  outward  and  forward,  between 
the  area  postrema  and  the  ala  cinerea. 

The  fovea  superior  is  accompanied  laterally  by  a  bluish  colored  area,  the  locus  caeru- 
leus.  The  latter  and  the  superior  fovea  exhibit  small  furrows  and  folds,  rugae  loci 
caerulei  et  foveae  superioris,  which  may  often  be  followed,  for  a  considerable  distance,  for- 
ward tpward  the  isthmus  and  backward  toward  the  recessus  lateralis.  To  the  outer  side 
of  the  sulcus  limitans,  lateral  to  the  fovea  superior,  the  fovea  inferior  and  the  ala  cinerea, 
the  area  acustica  is  seen  as  a  flat  elevation,  which  toward  the  recessus  lateralis  presents 
the  tuberculum  acusticum.  The  funiculus  separans,  above  noted,  courses  toward  the 
lower  inner  end  of  the  area  acustica  and  there  disappears. 


THE  GRAY  MASSES  OF  THE  RHOMBENCEPHALON. 

In  the  floor  of  the  trigonum  lemnisci,   in  the  isthmus,  lies  the  nucleus  lemnisci. 

The  pons  includes  a  larger  ventral  portion,  the  pars  basilaris  pontis,  and  a  smaller 
dorsal  one,  the  pars  dorsalis  pontis.  These  two  divisions  are  readily  seen  in  a  cross- 
section.  The  basal  part  exhibits  numerous  transversely  coursing  white  fibre-strands  that 
continue  laterally  into  the  pontile  crura  or  middle  cerebellar  peduncles.  In  the  lower 
part  of  the  basilar  division,  between  the  thin  white  fibre-bundles,  grayish  lamellae  repre- 
sent the  cross-sections  of  the  tracts  of  fibres,  which  descend  from  the  cerebral  peduncles, 
pass  through  the  entire  pons  and  continue  to  the  medulla  oblongata  and  the  spinal  cord. 
These  are  the  pyramidal  tracts,  the  fasciculi  longitudinal es  pyramidales.  The  fibrae 
pontis  superficiales  are  seen  as  transversely  coursing  fibres  that  pass  ventral  to  the  pyram- 
idal tracts,  while  the  fibrae  pontis  profundae  run  dorsal  to  or  partly  through  the 
pyramidal  strands.  The  pontile  nuclei,  nuclei  pontis,  are  small  masses  of  gray  substance 
lying  scattered  between  the  bundles  of  fibres. 


nervi  cochleae, 


NUCLEI    OF  THE   HIND-BRAIN.  83 

The   pars    dorsalis    pontis,   also   termed  the  tegmentum  pontis,   appears  gray  in 
transverse  sections.      It  contains  the  following  nuclei : — 

The  nucleus  nervi  abducentis,  within  the  colliculus  facialis, 

The  nucleus  nervi  facialis, 

The  nucleus  motorius  et  sensibilis  nervi  trigemini, 

The  nucleus  tractus  spinalis  nervi  trigemini, 

The  nuclei  nervi  acustici,  within  the  area  acustica,   embracing  : — 

Nucleus  medialis 

Nucleus   dorsalis 

Nucleus  medialis 

Nucleus  lateralis  (Deiters) 

Nucleus  superior  (Bechterew) 

Nucleus  n.   vestibularis  spinalis 
The  nucleus  olivaris  superior, 
The  nucleus  corporis  trapezoidei, 
The  nuclei  reticulares  tegmenti. 

Within  the  cerebellum  (Fig.   87),  in  addition  to  the  cortex  or  substantia  corti- 
calis    covering  the   entire    surface,    special    gray    masses    are    found    within    the    corpus 


nervi  vestibuli, 


Tegmentum  pontis. ._. 


N.  trigeminus 


.   Ventricle. 


_-____.  Motor  and  sensory 
jSttj.          trigeminus  nuclei 


Stratum 
profundum 
pontis 


Pyramidal  tracts 
\ Stratum  superfciale  pontis 


FIG.  86. — Cross-section  of  brain-stem  in  region  of  pons. 


medullare.  In  the  medial  part  of  the  hemisphere  lies  the  nucleus  dentatus, 
which  appears  as  a  much  plicated  lamella  of  gray  substance  with  a  medially  directed 
opening,  the  hilus  nuclei  dentati.  Within  the  worm,  the  roof -nucleus ,'  nucleus  fastigii 
or  nucleus  tecti,  lies  on  each  side  of  the  mid-line.  Between  the  nucleus  fastigii  and  the 
nucleus  dentatus,  two  additional  centres  are  found,  the  nuclei  globosi,  small  gray 
masses  lateral  to  the  roof-nucleus,  and  the  nucleus  emboliformis ,  medial  to  the 
dentate  nucleus. 


84 


MORPHOLOGY. 


Nucleus  fastigii 


Nuclei  globosi 
Nucl.  emboliformis 


Nucleus  dentahis 


Within  the  medulla   oblongata,  in   the  fasciculus  gracilis  within  the  clava,  lies 
the  nucleus  fasciculi  gracilis,  while  in  the  fasciculus  cuneatus,  in   its  position  correspond- 

ing to  the  tuberculum  cuneatum, 
lies  the  nucleus  fasciculi  cuneati. 
The  tuberculum  cinereum  corre- 
sponds to  the  nucleus  tractus  spi- 
nalis  nervi  trigemini.  Within  the 
olive  are  found  the  micleus  oliva- 
ris  inferior,  with  the  nuclei  of  the 
two  accessory  olives,  the  nucleus 
olivaris  accessorius  ventralis  and 
dorsalis.  The  nuclei  arcuati  lie 
ventral  to  the  pyramidal  tracts, 
while  within  the  lateral  columns 

he     nuclei 


PIG.  87.  —  Horizontal  section  of  the  cerebellum,  exposing  the  internal  nuclei 


FIG.  88. — Diagram  of   the  brain-stem,  dorsal   aspect,  showing   the  location  of   the   nuclei  of   the  cerebral  nerves.      Motor 
nuclei  are  red,  sensory  nuclei  are  blue. 

The  floor  of  the  trigonum  hypoglossi  contains  the  nucleus  nervi  hypoglossi.     Close 
to  the  latter  but  within  the  floor  of  the  ala  cinerea,   is  the  sensory  nucleus  of  the  vagus 


SUMMARY   OF    RHOMBENCEPHALON.  85 

nerve,  which  anteriorly  is  continuous  with  the  like  nucleus  of  the  glossopharyngeal  nerve. 
In  this  region,  medial  to  the  ala  cinerea,  the  motor  nucleus  dorsalis  of  the  glossopharyn- 
geal and  vagus  nerves  appears  as  a  small  group  of  cells.  The  nucleus  tractus  solitarii 
occupies  the  elongation  of  the  sensory  nucleus  of  the  glossopharyngeal  and  vagus  nerves. 
Somewhat  lateral,  but  more  deeply  placed,  lies  the  nucleus  ventralis  or  nucleus  ambiguus 
of  the  ninth  and  tenth  nerves.  The  caudal  prolongation  of  the  nucleus  ambiguus  con- 


FIG.  89. -.-Diagram   of  brain-stem,  lateral   aspect,  showing  location   of   nuclei   of  cerebral  nerves.      Motor  nuclei  are  red, 

sensory  nuclei  are  blue. 

tains  the  elongated  nucleus  nervi  accessorii,  whose  spinal  part  reaches  into  the  ventral 
horn  of  the  spinal  cord.  The  nerve-cells  within  ti\eformatio  reticularis,  occurring  scattered 
or  in  small  groups,  constitute  the  nucleus  of  the  formatio  reticularis. 

The  more  important  of  these  nuclei  are  discussed  at  greater  length  in  connection 
with  the  Fibre-Tracts.  The  positions  of  the  nuclei  of  the  cerebral  nerves  are  diagram- 
matically  shown  in  Figs,  88  and  89. 


SUMMARY   OF   THE   RHOMBENCEPHALON. 
To  the  rhombencephalon  or  hind-brain  belong : 

The  isthmus  rhombencephali, 
The  metencephalon, 
The  myelencephalon. 
It  encloses  the  fourth  ventricle. 

To  the  isthmus  rhombencephali  belong  : 

Dorsal — The  brachia  conjunctiva  cerebelli, 
The  velum  medullare  anterius, 
The  trigonum  lemnisci  ; 
Ventral — The  crura  cerebri. 

To  the  metencephalon  belong  : 

The  pons  and  the  cerebellum. 


86  MORPHOLOGY. 

The  cerebellum  is  subdivided  into  the  worm  and  the  hemispheres.  More  or  less 
deeply  penetrating  fissures  separate  the  lobes  of  the  hemispheres  from  one  another.  The 
chief  segments  are  the  lobus  superior,  the  lobus  posterior  and  the  lobus  inferior,  each  of 
which  is  made  up  of  lobules.  The  individual  lobes  and  lobules  of  the  hemispheres  corre- 
spond to  definite  divisions  of  the  worm. 

The  myelencephalon,  or  the  medulla  oblongata,  has  as  its  upper  boundary, 
ventrally  the  lower  border  of  the  pons,  dorsally  the  striae  medullares  fossae  rhomboideae. 
Below,  the  medulla  passes  into  the  spinal  cord,  its  ventral  boundary  being  the  lower  end 
of  the  pyramidal  decussation.  Dorsal,  behind  the  rhomboid  fossa,  are  the  dorsal  and 
lateral  columns,  with  their  tubercula,  and  the  restiform  bodies.  Ventral,  lie  the  pyramids 
and  the  olives. 

The  fourth  ventricle  has  as  its  roof  the  velum  medullare  anterius,  the  brachia 
conjunctiva  cerebelli,  the  velum  medullare  posterius  and  the  tela  chorioidea  ;  as  its  floor, 
the  fossa  rhomboidea.  It  is  connected  with  the  third  ventricle  by  means  of  the  aquae- 
ductus  cerebri,  below  is  continuous  with  the  central  canal  of  the  spinal  cord,  and  com- 
municates with  the  subarachnoid  space  by  means  of  the  apertura  mediana  (foramen 
Magendii)  and  the  aperturae  laterales  (foramina  Luschkae). 

The  most  important  masses  of  gray  substance  within  the  rhombencephalon  are  : 

The  nucleus  lemnisci, 

The  nucleus  ponds, 

The  substantia  corticalis  cerebelli, 

The  nucleus  dentatus 

The  nucleus  fastigii 

™,  ,  i  u     •  cerebelli, 

Ine  nucleus  globosi 

The  nucleus  emboliformis 

The  nucleus  gracilis, 

The  nucleus  cuneatus, 

The  nucleus  lateralis, 

The  nucleus  arcuatus, 

The  nucleus  olivaris  inferioris, 

The  nuclei  nervorum,  within  the  floor  of  the  fourth  ventricle. 


THE   MENINGES. 

The  membranes  investing  the  brain  are  three  :  the  dura  mater,  the  arachnoid  and 
the  pia  mater. 

The  dura  mater  forms  the  outermost  covering  of  the  brain.  Beneath  it  lies  the 
arachnoid,  a  delicate  transparent  membrane  that  is  separated  from  the  dura  by  the  sub- 
dural  space.  The  innermost  covering  is  the  pia  mater,  separated  from  the  arachnoid  by 
the  subarachnoidal  space.  The  arachnoid  and  the  pia  have  been  also  regarded  as  the 
outer  and  inner  layers  of  the  soft-brain  membrane,  the  leptomeninx,  in  contrast  to  the 
hard  brain-membrane,  the  pachymeninx,  represented  by  the  dura. 


BRAIN-MEMBRANES. 


DURA    MATER. 

The  dura  mater  consists  of  two  lamellae.  The  outer  lamella,  which  lies  against  the 
bone  and  serves  as  the  inner  periosteum  of  the  cranial  case,  consists  of  soft,  loose  vascular 
connective  tissue.  The  inner  lamella  is  denser,  made  up  of  fibrous  connective  tissue,  and 
contains  few  blood-vessels.  While  the  outer  layer  appears  as  periosteum  and  is  prolonged 
on  the  cerebral  nerves  as  robust  sheaths,  the  inner  layer  comes  into  closer  relation  with  the 
brain,  since  it  sends  processes  between  the  larger  divisions  of  the  brain.  Such  processes  are  : 

i.  The  falx  cerebri,  or  falx  cerebri  major,  which,  penetrating  between  the  hemi- 
spheres, begins  in  front  at  the  crista  galli,  attached  by  its  convex  upper  border  to  the 
sides  of  the  sulcus  sagittalis  of  the  cranial  vault,  and  extends  backward  as  far  as  the 
protuberantia  occipitalis  interna.  Between  the  outer  and  inner  lamellae,  along  the  upper 
convex  border  of  the  falx,  is  a  blood-space,  which  is  triangular  in  cross-section  and  known  as 
the  sinus  sagittalis  superior.  The  lower  concave  border  of  the  sickle-like  falx  is  free  and 
encloses  the  smaller  sinus  sagittalis  inferior.  From  the  internal  occipital  protuberance 

Sinus  sagittalis  superior     Granulationes  Pacchioni 


Cranium 

Spatium  epidnral. 

Dura  mater 

Spatium  subdural. 

A        h.       ' fi 
Spatium  subarach. 
Pia  mater 


Substantia  corticalis  cereb 


FIG.  90. — Schematic  section  through  the  skull  and  the  meninges. 

forward,  the  falx  is  attached  to  the  tent-like  tentorium  cerebelli,  the  line  of  junction  being  the 
tent-edge,  while  the  border  attached  to  the  crista  galli  is  the  crest-edge.  In  front,  the  falx 
only  incompletely  separates  the  two  frontal  lobes,  but  behind  its  height  is  so  increased, 
that  it  penetrates  almost,  but  not  quite,  to  the  upper  surface  of  the  corpus  callosum. 

2.  The  falx   cerebelli,   or  falx  cerebri  minor,  which   forms  a    small  sagittal   pro- 
longation of    the  large  falx,  penetrates  between   the  cerebellar  hemispheres  and  descends 
from  the   internal    occipital    protuberance  to  the  foramen    magnum.      The  convex  border 
encloses  the  sinus   occipitalis  and  is  attached  along  the  crista  occipitalis.     Corresponding 
to  the  terminal  limits  of    the  crest,  the  cerebellar  falx  divides  into   two  diverging  arms, 
which  enclose  the  continuations  of   the  sinus  occipitalis. 

3.  The    tentorium    cerebelli,   which   forms   a  dorsally  arched  transverse  partition 
between  the  basal  surface  of  the  occipital  lobes  of  the  cerebrum  and  the  dorsal  surface  of 
the  cerebellum.       The  outer  -convex   margin    is   attached,  on   each  side,  along  the  lineae 
transversae  of   the  occipital   and  'parietal   bones,  where  it  encloses  the  sinus  transversus, 
and  along  the  dorsal  edge  of  the  petrous  portion  of  the  temporal  bone,  where  it  conveys 
the  sinus  petroszis  superior.     Thence  the  attachment  of  the  tentorium   passes  to  the  pro- 


88  MORPHOLOGY. 

cessus  clinoides  anterior.  In  front,  the  free  inner  margin  of  the  tentorium  meets  the  outer 
one  and  then  extends  backward  and  slightly  upward  to  unite  with  the  lower  edge  of  the 
falx  cerebri.  Along  this  line  of  union  lies  the  sinus  rectus,  which  in  front  receives  the 
vena  cerebri  magna  Galeni  (Fig.  55)  and  behind  opens  into  the  confluens  sinuum  or  the 
torcular  Herophili. 

4.  The  diaphragma  sellae  turcicae,  which  forms  a  bridge  of  dural  tissue  over 
this  depression  in  the  sphenoid  bone.  Between  the  basal  and  dorsal  lamellae  of  the 
diaphragma  sellae  turcicae  lies  the  hypophysis  or  pituitary  body.  An  opening  in  the  middle 
of  the  membrane,  the  foramen  diaphragmatis ,  affords  passage  to  the  infundibulum. 

ARACHNOIDEA. 

This  delicate  transparent  membrane  consists  of  connective  tissue  and  is  devoid  of 
blood-vessels.  It  is  separated  from  the  dura  by  the  subdural  space  and  connected  with 
the  pia  by  strands  of  connective  tissue.  These  strands  form  the  subarachnoidal  tissue, 
and  the  cleft  between  the  arachnoid  and  the  pia  is  the  subarachnoidal  space.  The  latter 
is  traversed  by  the  connective  tissue  trabeculse  and  plates  of  the  subarachnoidal  tissue 
and  contains  a  fluid,  the  liquor  cerebro-spinalis,  in  considerable  quantity.  The  subarach- 
noidal space  communicates  with  the  ventricles  by  means  of  the  foramen  Magendii  and 
the  foramina  Luschkae  (page  80).  Over  the  cerebral  convolutions  the  subarachnoidal 
tissue  is  scanty,  in  these  localities  the  arachnoid  and  the  pia  being  fused  into  a  common 
membrane.  Over  the  cerebral  fissures,  on  the  contrary,  the  space  between  the  two 
membranes  is  larger,  since  the  pia  penetrates  into  the  fissures.  The  large  spaces  are 
found  principally  at  the  base  of  the  brain  and  where  the  latter  passes  into  the  spinal 
cord  ;  in  these  locations,  at  certain  places,  the  arachnoid  is  widely  separated  from  the  pia, 
resulting  in  the  formation  of  the  .cisternae  subarachnoideales.  Such  spaces  are  : 

The  cisterna  cerebello-medullaris,  between  the  posterior  border  of  the  cerebellum  and 
the  medulla  oblongata  ; 

The  cisterna  fossae  Syh'ii,   over  the  Sylvian  fossa  ; 

The  cisterna  chiasmatis,   surrounding  the  optic  chiasm  ; 

The  cisterna  interpeduncularis ,   between  the  cerebral  crura  ; 

The  cisterna  ambiens,  ascending  laterally  from  the  cerebral  peduncles  to  the  corpora 
quadrigemina  ; 

The  cisterna  corporis  callosi,  along  the  convex  dorsal  surface  of  the  callosum. 

In  certain  places,  as  on  both  sides  of  the  sinus  sagittalis  superior  or  along  the  sinus 
transversus,  villous  projections  from  the  outer  surface  of  the  arachnoid  push  before  them 
the  thin  dura  mater  and  encroach  on  the  venous  sinuses.  Such  elevations  are  called  the 
arachnoidal  villi  or  Pacchonian  granulations  (Fig.  90).  According  -to  the  investigations  of 
Key  and  Retzius,  these  structures  facilitate  the  passage  of  serous  fluid  into  the  venous  spaces. 

PIA   MATER. 

The  innermost  brain-membrane  consists  of  delicate  bundles  of  connective  tissue,  con- 
tains numerous  blood-vessels,  and  directly  invests  the  surface  of  the  brain,  penetrating  to 
the  bottom  of  all  the  fissures.  By  means  of  the  subarachnoidal  tissue,  the  pia  is  attached 
to  the  arachnoid.  Between  the  pia  and  the  surface  of  the  brain,  there  exists  only  a  very 
narrow  cleft,  the  subpial  or  epicerebral  space. 


THE   SPINAL    CORD. 


89 


THE  SPINAL  CORD: 

The  spinal  cord  or  medulla  spinalis  presents  a  compressed  cylindrical  column,  some- 
what more  flattened  in  front  than  behind,  that  is  enclosed  within  its  protecting  membranes 
and  only  incompletely  fills  the  vertebral  canal.  Above,  it 
passes  into  the  medulla  oblongata,  the  upper  limit  corre- 
sponding to  the  lower  end  of  the  pyramidal  decussation. 
Below,  the  spinal  cord  reaches  to  the  level  of  the  first 
or  second  lumbar  vertebra.  It  is  not  everywhere  of 
equal  thickness,  but  in  two  places  exhibits  spindle-shaped 
enlargements  (Fig.  91)  : 

a.  In    the    cervical    region    of    the  spine  the  cervical 
enlargement,  intiimescentia  cervicalis,  from  the  third  cervical 
to  the  second  thoracic  vertebra  ; 

b.  In  the  lower  part  of  the  thoracic  spine  the  lumbar 
enlargement,  intumescentia  lumbalis,  from  the  ninth  thoracic 
to  the  second  lumbar  vertebra. 

Both  enlargements  correspond  to  the  regions  in  which 
the  large  limb-nerves  arise. 

The  lumbar  enlargement  below  passes  over  into  a 
short  conical  segment,  the  conus  medullaris  or  conus 
terminalis,  from  which  proceeds  a  long  delicate  thread-like 
process,  the  filum  terminale. 

The  average  length  of    the    spinal  cord  is  45    cm.   in 
men,  and  from  41-42  cm.   in  women.       In  accordance  with 
the  pairs  of  spinal  nerves  given  off  from  the  cord,  we  recog- 
nize a  pars  cervicalis,  from  which  the  cervical  nerves  emerge  ;  a  pars  thoracalis,  from  which 
the  thoracic  nerves  arise,  and  a  pars  lumbalis,  from  which  the  lumbar  and  the  sacral  nerves 
are  derived. 

EXTERNAL   CONFIGURATION. 

The  anterior  or  ventral  surface  of  the  spinal  cord  is  cleft  in  the  mid-line  by  a  deep 
longitudinal  furrow,  \\\o.fissura  mediana  anterior ;  the  posterior  or  dorsal  surface  is  modelled 

by  a   superficial   longitu- 

viermedius  posterior  ^ina!    grOOVC,    the     SUlcuS 

medianus  posterior.  By 
means  of  these  two  fur- 
rows the  spinal  cord  is 
divided,  into  symmetrical 
halves.  Lateral  to  the 
posterior  median  sulcus, 
in  each  half  runs  the 
sulcus  lateralis  posterior, 
along  which  the  pos- 
terior root-bundles  enter. 


Intumescent. 
lumbalis 


Conus 
edullarit 


Filum 
terminals 


FIG.    91. — Front   view    of   the  spinal 
cord.    Schematic. 


Sulc.  med. 

post.        Sulcus 


Funicnlus  late; 


Sulcus  lateralis  posterior 
Cornu  posteriits 
Formatio  rettcularis 
Colutnna  lateralis 


ediana  anterior 


Radix  anterior 

Funicul, 
FIG.  92. — Transverse  section  of  the  spinal  cord. 


Cornu  antenus 
S,tlc.  lat.  anterior 


MORPHOLOGY. 


Lateral  to  the  anterior  median  fissure,  on  each  side  extends  the  sulcus  lateralis 
anterior,  which  is  not  a  continuous  furrow,  unless  the  emerging  anterior  root-fibres  are 
torn  away.  In  the  upper  thoracic  and  the  cervical  region,  an  additional  delicate  longi- 
tudinal groove,  the  sulcus  intermedius  posterior,  is  distinguishable  between  the  median 
and  lateral  posterior  sulci.  The  anterior  root-fibres  that  emerge  along  the  anterior  lateral 

sulcus  form  individual  bundles, 
the  radices  anteriores,  sepa- 
rated from  one  another  by 
intervals.  The  posterior  root- 
fibres,  which  enter  along  the 
posterior  lateral  sulcus  in  an 
unbroken  row,  likewise  form 
outwardly  converging  bundles, 
the  radices  posteriores.  Each 
pair  of  anterior  and  posterior 
Kadi*  ant.  root-bundles  passes  to  a  defi- 

FIG.  93. — Schematic   representation    of    the  formation  of  the   spinal  nerves.        nite       foramen 


Radix  Post. 


Ganglion  spin 


(Fig-  93)-  Here,  the  posterior 

root  presents  a  small  fusiform  swelling,  the  ganglion  spinale,  and  then  unites  in  its  fur- 
ther course  with  the  corresponding  anterior  root,  thereby  forming  the  spinal  nerve, 
which  latter  divides  into  an  anterior  and  posterior  division. 

The  emerging  root-bundles  run  not  only  outward,  but  at  the  same  time  caudalward, 
and,  indeed,  the  more  so  the  nearer  to  the  caudal  end  of  the  spinal  cord  they  emerge. 
In  the  lumbar  region,  the  course  of  the  nerve-roots  within  the  vertebral  canal  is  nearly 
parallel  with  the  long  axis  of  the  cord,  so  that  the  conus  medullaris  and  the  filum 
terminale  lie  in  the  midst  of  a  generous  bundle  of  nerve-roots,  which,  on  account  of  the 
supposed  resemblance  to  a  horse's  tail,  is  designated  the  cauda  equina. 

By  means  of  the  longitudinal  furrows,  the  spinal  cord  is  subdivided  into  the  following 
columns  : 

The  funiculus  anterior,  between  the  anterior  median  fissure  and  the  anterior  lateral 
sulcus  ; 

The  funiculus  lateralis,  between  the  anterior  and   posterior  lateral  sulci  ; 

The  fumculus  posterior,  between  the  posterior  median  fissure  and  the  posterior 
lateral  sulcus.  The  posterior  column  is  separated  by  the  sulcus  intermedius  posterior 
into  a  medial  and  a  lateral  division,  the  medial  one  being  known  as  the  fasciculus 
gracilis,  or  Gall's  column,  and  the  lateral  one  as  the  fasciculus  cuneatus,  or  Burdock's 
column. 

INTERNAL   CONFIGURATION. 

Even  with  the  unaided  eye,  one  can  readily  distinguish  gray  and  white  substance 
in  a  transverse  section  of  the  spinal  cord.  When  cut  across,  the  centrally  situated  gray 
substance  appears  H-form  in  outline.  The  bridge  of  gray  substance  connecting  the  two 
limbs  of  the  H,  encloses  the  central  canal,  canalis  centralis,  which  is  immediately  sur- 
rounded by  the  substantia  gelatinosa  centralis  and  lined  with  ependyma.  Above,  the 
central  canal  widens  at  the  transition  of  the  spinal  cord  into  the  medulla  oblongata  and 


THE  SPINAL   CORD.  91 

passes  over  into  the  fourth  ventricle.  Below,  at  the  lower  end  of  the  conus  terminalis, 
it  expands  into  the  ventriculus  terminalis  (Krause),  becomes  again  narrow  at  the 
transition  into  the  filum  terminale  and,  finally,  ends  blindly. 

The  part  of  the  gray  bridge  that  passes  behind  the  central  canal  is  known  as  the 
commissura  posterior,  that  which  lies  in  front  is  the  commissura  grisea  anterior.  In 
front  of  the  latter,  between  it  and  the  bottom  of  the  anterior  median  fissure,  is  the 
commissura  alba  anterior. 

The  gray  substance,  in  each  half -of  the  spinal  cord,  presents  in  front  a  thick 
swelling,  the  anterior  horn  or  cornu  anterius,  and  behind  a  more  slender  part,  the  poste- 
rior horn  or  cornu  posterius.  Since  the  gray  substance  extends  continuously  throughout 
the  entire  length  of  the  cord,  the  anterior  and  posterior  horns  appear  in  longitudinal 
sections  as  columns  ;  they  are  called  also,  therefore,  the  columnae  griseae.  The  lateral 


Lissaner's  zone 


Posterior  root 


Fortnatio  reticularis 


Clarke's  column 


Anterior  horn 


Commissura  alba  Anterior  root 

FIG.  94. — Transverse  section  of  the  spinal  cord. 

part  of  the  gray  substance,  in  the  lower  cervical  and  the  upper  thoracic  regions  of  the 
cord,  becomes  more  independent  and  there  forms  the  lateral  horn  or  columna  lateralis. 
In  the  entire  cervical  and  upper  thoracic  cord,  the  gray  substance  extends  into  the  white 
matter  as  a  network  of  gray  trabeculae  and  strands,  which  occupy  the  angle  between 
the  lateral  and  posterior  horns  and  constitute  the  formatio  reticularis.  The  posterior 
cornu  begins  ventrally  as  the  base,  then  becomes  narrower  and  forms  the  neck,  cervix 
columnae  posterioris ;  dorsally  follow  the  head  of  the  horn,  caput  columnae,  and  the 
point,  apex  columnae  posterioris,  which  latter  embraces  a  crescentic  field,  the  substantia 
gelatinosa  Rolandi,  and  the  dorsally  situated  marginal  zone.  Medial  to  the  neck  of  the 
posterior  cornu  and  close  to  the  posterior  commissure,  one  finds  the  nucleus  dorsalis  or 
Clarke 's  column  as  a  small  group  of  cells  within  the  gray  substance  of  the  upper 
lumbar,  the  entire  thoracic  and  the  lower  cervical  regions. 

The  white  substance  surrounds  the  gray  and  is  subdivided,  as  already  noted, 
into  three  tracts — the  anterior  column,  between  the  anterior  median  fissure  and  the  anterior 
roots,  the  posterior  r.olumn,  between  the  posterior  median  fissure  and  the  posterior  roots, 


92 


MORPHOLOGY. 


and  the  lateral  column,  between  the  anterior  and  posterior  roots.  The  posterior  column 
is  further  divided  by  the  sulcus  intermedius  posterior  into  the  medially  situated  fasciculus 
gracilis  or  Goll's  column,  and  the  laterally  placed  fasciculus  cuneatus  or  Burdach's 
column. 

In  the  essentials,  the  make-up  of  the  spinal  cord  is  the  same  in  its  various 
segments,  the  central  gray  substance  in  the  characteristic  H  being  everywhere  enclosed 
by  the  white  matter.  The  size  and  form  of  the  cord  in  cross-sections,  as  well  as  the 
proportions  of  the  gray  and  white  substance,  however,  vary  in  the  individual  regions. 
In  regard  to  size,  the  stronger  development  in  the  cervical  and  lumbar  enlargements  is 
at  once  noticeable.  So  far  as  the  form  is  •  concerned,  transverse  sections  are  so  charac- 
teristic in  the  different  regions  that,  within  certain  limits,  the  region  from  which  a  section 
has  been  taken  can  be  determined  from  such  data  alone.  Thus,  cross-sections  of  the 
cord  in  the  cervical  region,  particularly  at  the  level  of  the  IV-VIII  nerve,  and  in  part 


TfcBT 


Li 


FIG.  95. — Transverse  sections  of  the  spinal  cord  at  different  nerve-levels.     C,  cervical;  Th,  thoracic;  L,  lumbar;  S,  sacral. 

also  in  the  highest  thoracic  segments  are  transversely  oval ;  in  the  thoracic  region  the 
cross-section  is  almost  circular ;  while  in  the  lumbar  region  it  is  more  quadrate,  with 
more  marked  ventral  flattening.  The  quadrate  form  is  especially  evident  in  the  sacral 
and  likewise  in  the  coccygeal  cord,  where,  however,  in  contrast  to  the  lumbar  region, 
the  strongest  flattening  is  dorsal  with  coincident  ventral  narrowing. 

Regarding  the  proportion  of  the  gray  and  the  white  substance,  it  is  readily  seen 
that  the  gray  substance  is  most  abundant  in  those  segments  from  which  the  large  limb- 
nerves  arise,  that  is  in  the  cervical  and  lumbar  enlargements.  In  these  segments  the 
great  development  of  the  anterior  horns  is  particularly  evident.  The  gray  substance  in 
the  dorsal  cord,  on  the  contrary,  is  poorly  developed,  the  H-form  being  here  seen  to 
best  advantage.  The  white  substance  exhibits  a  robust  development  in  the  cervical,  as 
'well  as  the  thoracic  region.  Towards  the  lumbar  cord  it  progressively  decreases  in 
amount  and,  in  the  sacral  region  and  toward  the  conus  medullaris,  the  white  matter  forms 
only  a  thin  peripheral  zone  surrounding  the  gray  matter,  which  at  these  levels  con- 
siderably exceeds  in  amount  the  white. 


SPINAL   MENINGES. 


93 


THE   MEMBRANES    OF   THE   SPINAL   CORD. 

As  is  the  brain,  so  also  the  spinal  cord  is  surrounded  by  three  envelopes — the 
dura  mater,  the  arachnoid  and  the  pia  mater. 

Dura  mater  spinalis.  This  membrane  forms  a  strong  fibrous  investment  consist- 
ing of  two  layers,  the  outer,  which  fuses  with  the  periosteum  of  the  vertebrae,  and  an 
inner,  which  is  the  spinal  dura  proper.  The  space  between  these  two  layers  is  filled 
with  loose  connective  tissue,  contains  the  large  venous  plexus  and  is  traversed  by  lymph- 
spaces  ;  it  is  the  cavum  interdurale  or  cavum  epidurale.  The  dura  extends  as  a  long 


•^exf'. 

M^^ 


/4?^£%i^Q 


•^G^ZZ^-^  ir&lVWBjS^ 
fe^T^^lPS*** 


FIG.  96. — Schematic    representation   showing   relations  of  the  spinal  meninges  to  one  another  and  to  the  vertebral  canai. 
Dura  is  yellow;  arachnoid,  green;  pia,  with  ligamentum  denticulatum,  blue.  . 

wide  sac  over  the  conus  medullaris,  narrows  at  the  level  of  the  second  or  third  sacral 
vertebrae,  thence,  as  the  filum  durae  matris  spinalis,  clothes  the  filum  terminale  and 
finally  passes  into  the  periosteum  of  the  coccyx. 

Arachnoidea  spinalis.  This  is  a  delicate  vascularless  membrane,  separated  from 
the  dura  mater  by  the  cavum  subdurale  and  from  the  pia  mater  by  the  cavum  subarachnoi- 
deale.  It  is  connected  with  the  subarachnoidal  fibres,  which  are  particulary  robust  and 
numerous  toward  the  sulcus  medianus  posterior  ;  in  the  lower  cervical  and  in  the  thoracic 
region,  they  form  a  special  partition,  the  septum  subarachnoideale  or  septum  cevicale 
intermedum.  Within  the  subarachnoidal  space  the  liquor  cerebro-spinalis  circulates. 

Pia  mater  spinalis.  This  membrane  encloses  the  spinal  cord  as  a  delicate  vas- 
cular envelope  and  forms,  by  penetrating  within  the  anterior  median  fissure,  the  septum 
anterius.  The  pia  is  connected  with  the  dura  mater  by  means  of  the  ligamentum  dentic- 
ulatum. The  latter  consists  of  from  19-23  small  triangular  processes,  with  their  bases 
attached  to  the  pia,  which  extend  outward  from  the  lateral  surface  of  the  cord  between 
the  anterior  and  posterior  roots  of  the  spinal  nerves,  to  be  attached  by  their  points  to 
the  dura  mater.  The  ligamentum  denticulatum  serves  as  a  suspensory  band  that  holds 
the  spinal  cord  in  position. 


PART  II. 

THE  FIBRE-TRACTS. 


THE  FIBRE-TRACTS. 


METHODS   OF    STUDYING   THE   FIBRE-TRACTS. 

The  older  anatomists  contented  themselves  with  the  task  of  describing  the  brain 
simply  from  the  exterior  and,  in  a  sense,  without  further  leading  conceptions.  In  this 
period  originated  the  terminology  that  owes  its  existence  to  merely  purely  superficial  and 
accidental  resemblances.  As  examples,  one  recalls  the  designation  of  the  corpora  quadri- 
gemina  as  the  "nates"  and  "testes,"  the  suggested  resemblance  of  the  corpora  mamil- 
laria  to  the  female  breasts,  of  the  calcar  avis  to  the  cock's  spur,  of  the  lyra  Davidis  to 
a  harp,  or  of  the  fornix  to  an  arch. 

In  order  to  render  more  exact  investigation  possible,  the  pioneer  observers  first 
sought  to  overcome  the  softness  of  the  central  nervous  substance,  and  to  that  end  employed 
various  chemical  agents,  as  alcohol,  corrosive  sublimate  and  salt  solutions.  Cold  was 
also  used  to  give  the  brajn  greater  consistence,  and,  indeed,  Gennari  and  Reil  made  their 
observations  on  frozen  brains.  In  this  manner,  in  a  purely  morphological  way,  began  the 
foundation  of  the  study  of  the  internal  connection  of  the  individual  brain-segments  and. 
until  the  middle  of  the  nineteenth  century,  the  method  of  direct  mechanical  dissociation 
of  the  alcohol-hardened  brain  was  employed  to  demonstrate  the  chief  fibre-tracts  (Gall 
and  Spurzheim,  Burdach,  Reil,  Arnold,  Foville). 

A  distinct  advance  in  brain-anatomy  was  made  when  the  structure  of  the  central 
nervous  system  began  to  be  studied  from  the  standpoint  of  development.  In  this  Tiedemann 
and  Reichert  were  pioneers.  In  the  introduction  to  his  "Anatomic  und  Bildungsgeschichte 
des  Gehirns,"  Tiedemann  observes  that  the  origin  and  development  of  the  brain  were  an 
almost  totally  neglected  part  of  anatomy  and  physiology.  He  mentioned  the  law  formu- 
lated by  Harvey,  that  the  embryo  of  man  and  of  the  animals  does  not  appear  in  a  com- 
pleted and  only  diminished  form,  but  that  it  begins  with  a  simpler  form,  successively 
passes  through  lower  formative  stages  and  finally  reaches  a  higher  stage  of  development. 
Why,  says  Tiedemann,  should  not  a  similar  progression  from  a  simpler  to  a  more  complex 
structure  also  occur  in  the  construction  of  the  brain  of  the  embryo  and  of  the  foetus ; 
and,  further,  should  not  this  process  supply  explanations  concerning  the  form  and  struc- 
ture of  the  brain,  so  intricate  in  its  completed  condition?  Tiedemann  busied  himself  for 
several  years  with  the  construction  of  the  embryonal  and  foetal  brain.  The  pure  mor- 
phology of  the  brain,  however,  reached  high-water  mark  with  the  embryological  method 
of  examination  followed  by  C.  B.  Reichert.  Through  the  work  of  Schmidt,  Mihalkovics, 
Kolliker,  His  and  others,  this  method  has  led  to  a  strict  scientific  division  of  the  brain 
and  to  the  establishing  of  a  comprehensive  morphological  basis. 

7  97 


0S  THE   FIBRE-TRACTS. 

By  these  "  embryological "  methods  much  was  gained,  but  by  no  means  all. 
Embryology  taught  us  to  understand  the  development  of  the  form,  but  told  us  nothing 
concerning  the  intimate  connection  of  the  parts,  a  clear  insight  into  which  alone  leads  to 
a  comprehension  of  the  function  of  the  central  nervous  system.  The  question  of  the  intimate 
connection  of  the  parts,  however,  is  nothing  but  the  question  of  the  fibre-tracts  and  there- 
with we  enter  a  new  phase  of  brain-investigation.  We  may  designate  this  phase  as  the 
physiological  in  contrast  to  the  pure  morphological,  since  the  extraordinarily  difficult  and 
laborious  endeavors  of  the  later  investigators  to  unravel  the  intricate  fibre-complex  of  the 
central  nervous  system  are  all  undertaken  from  the  physiological  standpoint  and  with  a 
physiological  aim. 

After  Helmholtz  had  shown  for  the  invertebrates  and  Remak  for  the  vertebrates, 
that  the  nerve-fibres  proceed  from  the  nerve-cells,  it  became  evident  that  the  simple 
method  of  dissociation  no  longer  sufficed.  What  neurology  had  then  to  attempt  was 
not  merely  the  accurate  description  of  the  external  form,  but,  before  all  else,  the  estab- 
lishing and  the  tracing  of  the  intricate  paths  which  the  nerve-fibres  pursue,  and  the  definite 
establishment  of  all  the  numerous  connections  joining  centre  with  centre  within  the  interior 
of  the  central  nervous  system  and  bringing  the  latter  into  relation  with  the  periphery. 
Although  tracing  these  fibre-paths  even  within  the  peripheral  nerves  is  by  no  means  easy 
because  of  the  peculiar  plexus-formation  and  anastomoses  of  certain  nerves,  such  task  is 
especially  difficult  within  the  brain  and  spinal  cord,  since  here  often  within  a  small  space 
the  most  diverse  paths  run  side  by  side  and,  further,  decussations  and  interfeltings  of  the 
nerve-fibres  make  the  direct  tracing  of  the  nerve-tracts  impossible. 

A  method  of  fundamental  importance  for  following  the  nerve-paths  through  longer 
stretches  was  now  introduced,  namely,  the  method  by  series  of  consecutive  sections, 
introduced  by  Benedikt  Stilling.  The  necessity  of  cutting  the  brain  and  spinal  cord  into 
thin  segments  for  the  accurate  investigation  of  their  finer  structure,  even  the  older  inves- 
tigators recognized  and  suggested  means  by  which  to  accomplish  this  end.  As  early 
as  1824,  Rolando  made  thin  cross-sections  of  hardened  spinal  cord  with  a  razor  and 
examined  them  with  a  hand-lens.  But  the  segments  cut  by  Rolando  were  not  sufficiently 
thin  to  be  used  with  higher  amplification  ;  moreover,  they  were  made  without  system. 
In  1836  Valentin  examined  microscopically  the  spinal  cord  of  freshly-killed  sheep  and 
pigeons,  by  cutting  the  cord,  under  water,  with  a  pointed  two-bladed  knife,  into  the 
thinnest  possible  lamellae  and  then  carefully  compressing  the  sections  while  being  observed. 
In  this  manner  Valentin  studied  the  spinal  cord  layer  by  layer,  from  without  inward,  in 
longitudinal  sections,  and  expressed  the  opinion,  that  for  the  correct  understanding  of  the 
structure  of  the  spinal  cord  examination  by  strata  is  the  only  proper  way.  Four  years 
later,  Hannover  advanced  along  this  line  even  farther  than  Valentin.  He  employed  a 
brain  and  spinal  cord  hardened  in  chromic  acid,  which  he  cut  into  thin  sections  with  a 
sharp  knife,  and  examined  the  relations  of  these  lamellae  piece  by  piece. 

Shortly  after  the  appearance  of  Hannover's  paper,  the  great  doctor  of  Cassel, 
Benedikt  Stilling,  began  in  1841  his  investigations  concerning  the  structure  of  the  spinal 
cord.  Stilling  was  the  first  to  cut  a  spinal  cord  into  many  consecutive  sections,  as  thin 
and  transparent  as  possible,  and  then  to  study  in  each  section  the  distribution  of  the 
white  and  gray  substance.  He  traced  progressively  from  section  to  section  the  changes 
in  the  picture,  and  finally,  by  reproducing  the  individual  pictures,  gained,  at  least  to  a 


METHODS   OF  INVESTIGATION.  99 

certain  degree,  a  clear  conception  of  the  internal  structure  of  the  cord.  This  method  by 
series  of  consecutive  sections,  which  Stilling  designated  as  "investigation  layer  by  layer," 
is  even  to-day  the  one  most  employed  in  the  examination  of  the  central  nervous  system. 
During  the  continual  use  of  so  productive  a  method,  it  was  inevitable  that  the  original 
technique  of  Stilling  should  undergo  many  alterations  and  improvements.  The  employ- 
ment of  the  method  was  facilitated  by  better  hardening  of  the  organs.  Already  in  1832, 
Ludwig  Jacobson  recommended  potassium  chromate  as  a  preservative  for  anatomical 
preparations.  Hannover  first  put  Jacobson' s  discovery  to  use  for  histological  investigation. 
Later  chromic  acid  was  displaced  from  the  technique  by  one  of  its  salts.  At  any  rate, 
no  one  has  rendered  greater  service  than  Heinrich  Muller  by  the  introduction  of  potas- 
sium bichromate,  now  so  universally  serviceable.  From  him  came  also  the  classic  Muller' s 
fluid,  which,  indeed,  even  to-day  is  much  used  in  its  original  composition.  Later  followed 
many  new  hardening  reagents.  One  of  these,  formalin  or  formol,  must  be  especially 
mentioned,  since  in  recent  years  its  many  advantages  have  brought  it  into  universal  use. 
Formol  was  introduced  in  histological  technique  by  Blum  in  1893. 

The  method  of  consecutive  sections  was  further  facilitated  by  the  introduction  of  the 
microtome,  by  which  exact  cutting  and  large  regular  sections  are  made  possible,  so  that 
an  entire  brain  may  be  laid  into  a  series  of  thin  sections  without  losing  one.  We  may 
mark  the  sections  in  their  proper  sequence,  determine  in  each  the  topography  of  the 
gray  substance  and  of  the  fibre-tracts,  and,  by  means  of  the  series,  from  these  isolated 
data  construct  a  composite  picture  of  the  principles  of  construction  of  the  part  of  the 
brain  under  discussion. 

The  application  of  Stilling' s  mode  of  investigation  was  materially  facilitated  by  the 
methods  of  staining.  For  a  long  time  Gerlach's  carmine  staining  was  dominant..  An 
important  advance  was  gained  by  Weigert's  admirable  hematoxylin-method.  At  present 
we  have  at  our  disposal  a  quite  considerable  number  of  different  dyes,  which  may  be 
used  with  advantage  in  investigating  fibre-paths.  But  neither  the  Weigert  stain,  nor  any 
other  of  the  procedures  so  far  recommended  and  used,  is  capable  of  solving  the  question,  the 
answer  to  which  has  always  been  most  sought  for  the  correct  understanding  of  the  structure 
of  the  nervous  system.  Continually  was  asked  :  In  what  manner  are  the  nerve-fibres 
related  to  the  nerve-cells?  In  what  manner  are  the  nerve-cells  related  to  one  another? 
How  do  the  nerve-fibres  in  the  brain  and  spinal  cord  arise  and  how  do  they  end? 

In  this  connection,  two  methods  were  epoch-making — Ehrlich's  methylene  blue  method 
and  Golgi's  silver-method.  Ehrlich's  procedure,  which  was  introduced  in  1886  and 
depends  on  the  coloring  of  the  living  nerves  by  means  of  methylene  blue,  was  subse- 
quently improved  by  Retzius,  Apathy,  Bethe  and  others.  Golgi's  method  is  older.  A 
number  of  years  before,  the  Italian  investigator  had  obtained  preparations,  in  which  the 
nerve-cells  and  their  processes  stood  out  with  great  sharpness  as  dark  figures,  by  treating 
the  brain-substance  with  the  chromic  acid  salts  and  with  silver  nitrate.  Golgi  described 
his  method  as  early  as  1873,  but  at  first  his  observations  were  little  known.  Not  until 
the  publication  of  an  elaborate  paper  in  1886,  did  Golgi  excite  widespread  attention  and 
his  results  and  methods  become  the  starting  point  of  an  energetic  examination  of  the 
central  nervous  system.  For  example,  the  Spanish  savant,  Ram6n  y  Cajal,  was  able,  by 
the  use  of  the  Golgi  method  on  embryos  and  young  animals,  to  arrive  at  results  that 
partly  solved  many  of  the  dominant  questions,  or  placed  them  in  a  new  light.  At  first 


ioo  THE   FIBRE-TRACTS. 

through  the  labors  of  this  investigator,  soon  also  through  those  of  others,  especially  of 
Kolliker,  Lenhosse'k,  van  Gehuchten  and  Retzius,  a  clear  picture  took  the  place  of  the 
previous  schemata.  The  most  important  findings  of  these  researches  are,  that  the  nerve- 
fibres  are  nothing  more  than  extraordinarily  long  processes  of  the  nerve-cells,  that  every 
nerve-fibre,  from  beginning  to  end,  is  to  be  regarded  as  a  part  of  a  single  nerve-cell, 
and  that  every  nerve-cell,  with  the  nerve-fibres  proceeding  from  it,  represents  an  histo- 
logical  individuality  or  nervous  unit.  Waldeyer  christened  such  anatomical  unit,  neurone, 
and  therewith  founded  the  neurone-theory. 

The  method  of  Stilling  enables  us  to  trace  a  nerve-tract  through  a  long  stretch. 
The  identity,  however,  is  possible  and  certain  only  so  long  as  the  fibre-bundles  compos- 
ing the  tract  do  not  suffer  interruption,  or  so  long  as  they  are  not  deflected  from  the 
plane  of  the  section,  or  do  not  separate  into  widely  diverging  fibres.  The  accurate  iden- 
tification and  tracing  of  the  fibre-tracts,  even  when  they  branch  in  the  most  diverse 
directions  or  resolve,  have  necessitated  the  search  for  new  methods. 

One  of  these  additional  methods  is  the  pathologico- anatomical  one,  based  on  the  inves- 
tigation of  secondary  degenerations.  Rokitansky  announced  in  the  first  edition  of  his 
Pathological  Anatomy  (1847),  that  atrophy  of  the  brain  following  apoplexy  and  inflam- 
mation leads  to  atrophy  of  different  fibre-paths,  when  extensive,  indeed,  to  the  disap- 
pearance of  an  entire  hemisphere  and  the  related  fundamental  tracts.  This  communica- 
tion for  a  time  remained  unnoticed.  In  1850  Ludwig  Tiirck  described  more  closely  such 
secondary  degenerations  and  inferred  from  his  findings,  that  in  those  cases  of  cross-section 
of  the  spinal  cord,  in  which  the  direction  of  physiological  conductivity  and  that  of  the 
degeneration  were  identical  in  the  secondarily  degenerating  cord-paths,  the  degeneration 
itself  was  caused  by  the  disturbance  of  functions.  Notwithstanding  these  exceedingly 
important  results,  at  first  only  few  investigators  followed  Tiirck  along  this  line  of  investi- 
gation. In  later  years,  however,  this  method  has  been  universally  employed  and  to  it 
we  are  indebted  for  the  many  papers  by  which  our  knowledge  of  the  fibre-paths  of  the 
central  nervous  system  has  been  materially  extended.  The  method  depends  upon  the 
principle,  that  every  nerve-fibre  in  its  function  is  dependent  upon  the  related  nerve-cell. 
Destruction  of  the  latter,  or  separation  of  the  nerve-fibre  from  its  cell,  results  in  degenera- 
tion of  the  related  fibre.  Let  us  assume  that  a  descending  tract  of  the  spinal  cord  has 
been  destroyed  in  some  part  of  its  course.  What  happens  ?  The  portions  of  the  nerve- 
fibres  below  the  injury  are  separated  from  their  trophic  centre  ;  they  therefore  die.  This 
destruction  or  secondary  degeneration  within  the  spinal  cord  proceeds  downward.  On 
examining  a  cross-section  of  the  cord  passing  below  the  seat  of  injury  and  comparing  it  with 
a  corresponding  section  of  a  normal  spinal  cord,  the  seat  of  the  degeneration  is  readily 
located  and  the  involved  tracts  may  be  accurately  followed  by  means  of  serial  sections. 

This  method  of  investigation  by  secondary  degeneration  is  closely  related  to  the 
physiological  method. or  the  method  of  vivisection.  Certain  nerve-centres  or  nerve-fibres 
of  an  animal  may  be  directly  stimulated  or  destroyed,  and  from  the  resulting  symptoms 
conclusions  drawn  as  to  the  relations  of  the  nerve-centres  or  nerve-tracts  to  the  peripheral 
parts  ;  thereby  a  functional  differentiation  of  the  nerve-fibres  is  also  possible. 

The  pathological  method  is  based  on  a  principle  similar  to  that  of  the  vivisection 
procedures.  Here  also  the  destruction  of  parts  of  the  central  nervous  system  is  con- 
cerned, but  these  mutilations  are  not  experimental  but  caused  by  the  establishing  of 


METHODS   OF  INVESTIGATION.  101 

diseased  processes.  In  this  relation,  the  study  of  the  pathological  changes  in  certain 
affections  of  the  spinal  cord  is  of  primary  importance. 

By  means  of  the  experimental  method,  which  has  been  used  on  animals  with  such 
great  success,  we  are  able  to  follow  and  to  study  the  course  of  the  fibre-bundles  by 
degenerations.  This  method,  employed  only  under  certain  conditions,  was  introduced  by 
Gudden  and  his  pupils  and  is  the  atrophy-method,  or  the  method  of  developmental 
arrest.  Gudden' s  method  is  distinguished  from  other  experimental  procedures  in  that  it 
is  directed  against  the  young  animal.  The  chief  difference  consists  therein,  that,  follow- 
ing an  experimental  impression  on  the  new-born  animal,  the  entire  process  proceeds  much 
more  rapidly  and  completely  than  in  the  adult.  The  absorption  of  the  disintegration 
products  from  the  elementary  parts  destroyed  goes  on  much  more  rapidly  and  com- 
pletely in  the  new-born,,  so  that  scarcely  a  trace  of  the  fibres  and  only  few  remains  of 
the  cells  are  left.  In  addition,  .the  technique  is  relatively  easy,  while  a  further  distinct 
advantage,  as  Gudden  himself  pointed  out,  is  the  almost  incredibly  rapid  and  admirable 
healing  of  the  injury  without  disturbing  secondary  processes. 

In  1852  Waller  showed,  that  the  peripheral  stump  of  a  cross-sectioned  peripheral 
nerve  undergoes  degeneration.  For  a  long  time  it  was  believed,  that  the  peripheral 
segment  alone  degenerated,  and  that  the  central  one  remained  unaffected  by  such  changes. 
Since  the  study  of  Ranvier  on  degeneration  and  regeneration  of  sectioned  nerves,  how- 
ever, we  know  that  also  the  central  segment  suffers  important  modifications.  Ranvier 
showed,  that  in  the  central  segment  of  the  axis-cylinder  new  fibrillae  were  formed,  which 
became  new  nerves,  using  the  sheath  of  the  degenerating  peripheral  segment  as  a  sup- 
port to  reach  the  periphery.  The  nerve  reassumes  its  function — it  is  regenerated.  If, 
however,  from  any  cause  the  developing  nerve  fails  to  secure  such  support,  its  further 
development  is  arrested  and  a  nervous  tumor,  a  neuroma,  is  formed,  as  seen  in  amputation- 
stumps.  But  in  these  cases,  especially  when  of  long  standing,  a  certain  grade  of  atrophy 
of  the  nerves,  as  well  as  a  diminution  in  the  number  of  the  corresponding  nerve- 
cells,  may  be  observed.  These  changes  are  exceptionally  rapid  and  marked  so  soon 
as  the  interference  occurs  in  young  individuals,  particularly  in  the  new-born.  If  in 
a  new-born  animal  a  motor  nerve  is  removed,  a  certain  region  of  the  cerebral  cortex 
destroyed,  or  the  spinal  cord  partially  cut  through,  not  only  is  always  a  degeneration  of 
the  fibres  in  the  separated  peripheral  stump  to  be  observed,  but  also  atrophy  and  indeed 
complete  disappearance  of  the  cells  of  origin.  Gudden  believed  at  first,  that  this  differ- 
ence from  the  Wallerian  degeneration  was  attributable  to  the  lesion  being  in  the  new- 
born animal.  Later,  he  recognized  that  it  was  not  the  age,  but  the  position  that  exer- 
cised the  influence.  Then,  too,  Forel  proved,  that  the  death  of  the  cell  after  destruction 
of  the  related  fibre  occurred  in  the  adult,  as  well  as  in  the  new-born  animal.  Death  of 
the  cell  depends  alone  on  the  place  where  the  fibre  is  sectioned.  Section  of  a  motor 
nerve  at  the  periphery  is  followed  by  only  a  slow  impairment  and  diminution  in  the  size 
of  the  cells  and  fibres  of  the  central  stump.  Section  of  the  same  nerve  at  its  point  of 
emergence  from  the  brain,  results  in  the  death  of  the  central  root,  as  well  as  of  all  the 
cells  of  origin  within  the  nucleus  of  the  nerve.  The  method  of  Gudden  has  been  rich 
in  results.  In  1872-74  Gudden  proved,  by  extirpation  of  the  cortical  motor  zone  in 
dogs,  that  the  pyramidal  tracts  run  direct  from  the  cerebral  cortex  to  the  spinal  cord. 
Other  important  results  are  the  establishing  of  the  nuclei  of  origin  of  almost  all  the 


102 


THE   FIBRE-TRACTS. 


motor  cerebral  nerves,  the  course  of  the  medial  fillet  and  the  termination  of  the  optic 
tract.  Closely  connected  with  the  method  of  Gudden  are  the  pathological  cases  of  early 
lesion  and  consequent  atrophy  of  certain  parts  of  the  central  nervous  system,  as  well  as 
the  cases  of  congenital  malformation  involving  the  cerebro-spinal  axis. 

Our  kno\v!edge  concerning  the  fibre-tracts,  moreover,  has  been  especially  advanced 
by  the  embryological  method  introduced  by  Flechsig,  based  on  the  study  of  the  develop- 
ment of  the  nerve-fibres.  This  method  rests  on  the  fact,  that  the  different  fibre-systems 
within  the  central  nervous  system  acquire  the  medullary  substance  at  a  definite  time,  which, 
however,  varies  for  the  individual  systems.  On  examining  the  infantile  brain,  it  is  found 
that  certain  fibres  are  already  medullated,  while  others  have  not  yet  acquired  this  sheath. 
This  difference  between  the  medullated  and  non-medullated  fibres  is  readily  appreciable 
microscopically  and,  therefore,  the  examination  of  the  nervous  system  in  its  various  develop- 
mental stages  affords  the  possibility  of  delimiting  and  tracing  certain  fibre-systems. 

An  additional  means,  which  has  contributed  much  not  only  to  the  morphology  but 
also  especially  to  the  accurate  investigation  of  the  fibre-tracts,  is  the  comparative  anatom- 
ical method.  Since  in  the  different  classes  of  animals  this  or  that  part  of  the  brain  is 
varyingly  developed,  in  correspondence  with  different  functional  development,  the  investi- 
gations in  the  domain  of  comparative  anatomy  have  supplied  numerous  explanations  con- 
cerning the  many-sided  connections  of  individual  parts  of  the  central  nervous  system. 

Finally,  a  combination  of  these  various  methods  has  been  attempted.  Edinger 
united  the  comparative  anatomical  method  and  that  of  Flechsig.  Bechterew  combined 
vivisection  with  the  study  of  development  and  created  the  embryologico-physiological 
method.  Admirable  results  were  also  achieved  by  Bechterew  with  his  pathologico-physio- 
logical  method,  which  consisted  in  studying  secondary  degenerations  with  simultaneous 
stimulation  of  the  degenerated  parts  by  means  of  the  electrical  current. 

HISTOGENESIS  OF  THE  NERVOUS  SYSTEM. 

The  elements  of  the  nervous  system  are  developed  from  the  outer  germ-layer  or 
the  ectoderm.  As  we  have  already  seen,  the  brain  and  the  spinal  cord  arise  from  a  broad 
medial  strip  of  ectoderm.  Here  the  medullary  plate  is  formed,  which  is  bounded  exter- 
The medullary  plate  sinks  and,  at  the  same  time,  projects  with 
its  edges  above  the  level  of  the  embry- 
onic area  ;  in  this  way  is  formed  the 
medullary  groove,  bounded  by  the  med- 
ullary ridges.  The  medullary  groove 
closes  and  becomes  the  medullary  or 
neural  tube. 

The  medullary  tube,  the  direct 
successor  of  the  medullary  plate,  consists 
at  first  of  closely  pressed  epithelial  cells, 
each  of  which  extends  the  entire  thick- 

ness of  the  layer.  The  wall  of  the  entire  tube,  therefore,  originally  exhibits  the  character  of 
a  single-layered  columnar  epithelium,  whose  cells  are  bounded  by  the  membrana  limitans 
externa  on  the  one  side,  and  by  the  membrana  limitans  interna  on  the  other  (Fig.  97). 
Each  epithelial  cell  encloses  a  large  nucleus.  In  the  inner  zone,  other  large  cells  are  irregu- 


nally  by  the  cuticle-plate. 


FIG.  97. — Dorsal  half  of  neural  tube,  overlaid  by  the  ectoderm; 
epithelial  and  two  dividing  (so-called  germ-)  cells.     (His.) 


DEVELOPMENT  OF  NEUROGLTA.  103 

larly  scattered  between  the  epithelial  elements,  from  which  they  are  clearly  distinguished  by 
their  round  form  and  transparent  homogeneous  protoplasm.  His  designated  these  as  the 
germ-cells. 

The  epithelial  cells  multiply  rapidly  and,  consequently,  become  laterally  compressed 
and  elongated.  Their  nuclei  lie  at  different  heights  and  give  rise  to  the  appearance  of  a 
three-  to  six-celled  layer.  In  reality,  however,  the  cells  completely  retain  the  character 
of  a  single-layered  columnar  epithelium. 

Some  of  the  epithelial  cells  are  early  transformed.  They  grow  into  the  spongio- 
blasts  of  His,  from  which  are  developed  the  supporting  elements,  the  ependyma  and 
neuroglia  cells. 

Others  of  the  epithelial  cells  change  to  pear-shaped  elements  and  become  the  neuro- 
blasts,  which  are  transformed  into  the  nerve-cells. 

Both  kinds  of  cells,  the  spongioblasts  and  the  neuroblasts,  therefore,  are  derivatives 
from  the  original  ectodermic  elements  of  the  medullary  plate.  'The  above  mentioned 
"germ-cells"  of  His  are  nothing  more  than  cells  of  the  primary  medullary  area  under- 
going mitosis  and  represent  elements,  whose  division  supplies  the  material  for  the  increase 
of  the  indifferent  ectodermal  cells,  on  the  one  hand,  and  of  their  derivatives,  the  spongio- 
blasts and  the  neuroblasts,  on  the  other. 

DEVELOPMENT  OF  THE   EPENDYMA  AND  THE  NEUROGLIA  CELLS. 

The  ependyma  cells  maintain  in  the  foetal  stage  the  character  of  an  epithelium 
and  the  relations  to  the  membrana  limitans  externa  and  interna.  In  the  brain,  as  in  the 
spinal  cord,  the  ependyma  cells  extend  from  the  inner  to  the  outer  surface  of  the  neural 
wall,  the  length  of  the  cells  keeping  pace  with  the  increase  in  the  tube.  The  inner 
portion  of  the  cell,  nearer  the  central  canal,  retains  more  the  character  of  a  cell-body — 
ependyma  cell,  while  the  outer  portion  gradually  diminishes  to  a 
delicate  fibre,  which  as  an  ependyma  fibre  radially  traverses  the 
neural  wall.  The  entire  arrangement  constitutes  the  ependyma 
system  or  the  ependymium. 

On    examining   this   ependymal   framework    more    closely,  a 
distinctive  disposition  of   the  ependyma   cells  is  seen  in  the  spinal 

PIG.  98. — Transverse  sec- 

cord.  In  a  cross-section  of  the  medullary  tube  of  a  3-4  day  tion  of  neural  tube  of  a  four- 
chick  embryo  (Fig.  98),  we  recognize  how  the  ependyma  fibres  ^fecjlick  embryo-  (Len~ 
traverse  the  wall  of  the  tube,  at  the  sides  passing  almost  parallel 

from  the  central  canal  and  ventrally  and  dorsally  diverging  radially.  In  conse- 
quence of  the  coming  together  of  the  nucleus-bearing  portions  of  the  cells,  there 
appears  within  the  medullary  tube,  in  the  vicinity  of  the  central  canal,  a  broad, 
richly  nucleated  zone,  the  inner  layer  of  His  or  the  ependymal  nuclei-zone  of  Len- 
hosse"k.  In  a  general  way,  this  zone  corresponds  to  the  later  epitheiulm  of  the  central 
canal.  The  ependyma  fibres  of  the  later  anterior  commissure  present  a  rough  appearance, 
being  heavy  and  beset  with  spines  ;  they  also  emphasize  the  already  slight  meridional 
disposition  of  the  more  laterally  situated  ependyma  fibres.  In  a  somewhat  later  stage, 
the  ependyma  fibres  exhibit  varicosities,  particularly  in  their  inner  portions ;  in  addition, 
in  their  outer  portions  they  undergo  a  subdivision  into  several  branches,  all  of  wh'ch 
extend  to  the  periphery,  where  they  end  in  small  triangular  expansions. 


[04 


THE   FIBRE-TRACTS. 


Subsequent  stunting  of  the  lateral  parts  of  the  ependyma  framework  is 
most  marked  in  the  spinal  cord  of  the  higher  forms.  In  the  other  parts  of  the 
central  nervous  system,  the  ependyma  cells  and  fibres  retain  their  embryonal  form,  even 
after  completed  growth.  The  ependyma  cells  are,  therefore,  phylogenetically  as  well 
ontogenetically,  the  oldest  cells  of  the  supporting  framework,  arising  directly  from  the 

ectoderm  cells,  or,  indeed,  being  in  a 
modified  way  these  themselves.  During 
later  stages,  the  elements,  particularly 
the  ependyma  fibres,  are  curtailed  in 
varying  degrees  ;  a  part  of  the  epen- 
dyma cells  later  migrate  and  become 
the  neuroglia  cells. 

The  neuroglia  cells  arise  only 
after  the  formation  of  the  ependyma 
framework.  On  examining  the  spinal 
cord  of  a  ten-day  chick,  one  finds  a 
number  of  elements  which  closely  re- 
semble the  ependyma  cells,  their  fibres 
likewise  extending  to  the  periphery  and 
there  ending  in  conical  thickenings. 
They  differ  from  the  ependyma  cells 
proper,  however,  in  that  their  cell- 
bodies  no  longer  lie  at  the  central  canal,  but  farther  outward.  At  first  such  cells  are 
encountered  only  in  the  vicinity  of  the  central  canal  and  in  meagre  number ; 
later,  however,  they  are  more  numerous  and  occur  also  in  the  peripheral  zone. 
This  is  explained  by  the  manner  in  which  the  neuroglia  cells  arise.  Originally  these 
cells  lie,  as  do  the  ependyma  cells,  at  the  central  canal ;  then  the  cell-bodies  migrate 


FIG.  99. — Transverse  section  of  the  spinal  cord  of  a  human  embryo, 
14  cm.  long,  showing  ependymal  framework.     (Lenhossek.) 


FIG.  100. — Development  of  the  neu- 
roglia cells.  Spinal  cord  of  a  ten-day 
chick.  (Lenhossek.) 


FIG.  roi. — Xeuroglia  cells 
from  the  white  substance  of 
the  spinal  cord  of  an  embryo  of 
30  cm.  in  length.  (Lenhossek.) 


from  the  region  of  the  epithelium,  part  of  the  cell-body  becoming  a  thin  fibrilla,  that 
later  disappears.  Small  spines  and  branches  appear  on  the  former  smooth  cell-bodies, 
as  well  as  similar  thorny  outgrowths  for  a  short  distance  along  the  process  stretching 
from  the  cell-bodies  to  the  periphery.  At  first  such  migrated  cells  are  present  only  in 
sparing  number ;  later,  however,  their  number  materially  increases  and  the  cells  are  dis- 
tributed more  or  less  uniformly  throughout  the  entire  cross-section  of  the  spinal  cord. 


DEVELOPMENT  OF  NERVE-CELLS. 


105 


FIG.  102. — Transverse  sec- 
tion of  the  spinal  cord  of  a 
four-weeks  human  embryo. 
Differentiation  into  the  inner 
layer,  next  the  lumen  of  the 
canal,  the  middle  or  mantle 
layer,  containing  the  neuro- 
blasts,  and  the  outer  periph- 
eral layer.  (His.) 


This  radiating  sustentacular  apparatus  constitutes  in  man  and  the  higher  mammals  an 
embryonal  feature.  Subsequently,  the  picture  changes.  The  radial  type  disappears  and 
the  shape  of  the  cells  alters.  The  minute  spicules  and  branches  develop  very  markedly, 
while  the  peripherally  directed  processes  atrophy.  The  cells  become  the  true  spider  or 
neuroglia  cells.  The  latter,  therefore,  pass  through  various  developmental  stages ;  at 
first  they  are  ependyma  cells,  then  radial  sustenacular  cells,  from  which  arise  the  neurog- 
lia cells. 

DEVELOPMENT  OF  THE  NERVE-CELLS. 

The  neuroblasts,  from  which  the  nerve-cells  arise,  are  developed  in  the  innermost  layer 
of  the  medullary  tube,  bordering  the  central  canal.     Thence  they  migrate  outward  through 
the  inner  layer  and  localize  within  a  dorso-ventrally  expanding  region,  that  is  bounded  medially 
by  the  inner  layer  and  laterally  by  the  marginal  zone.      On  examining  a  cross-section  of  the 
medullary  tube  of  a  four-weeks  human  embryo  (Fig.  102),  the  cleft- 
like  central  canal  is  seen  in  the  middle,  bordered  by  the  inner  plate, 
outside  of  which  lies  the  stratum 
of  neuroblasts,    broad   ventrally 
and   thinner  dorsally.       Follow- 
ing His,  we  call  this  stratum  the 
mantle   layer.    ,  Peripheralward, 
the  latter  joins  the  marginal  zone 
— the  Randschleier  of  His. 

The  neuroblasts  are  pear- 
shaped  cells,  with  oval  nuclei, 
which  send  out  a  peripherally  di- 
rected process  that  bears  at  its 
end  a  characteristic  thickening, 

the  growth-wedge  of  Cajal.     This  process  is  nothing  less  than  the 
later  nerve-fibre.     While  the  rapidly  growing  fibres  endeavor  to 

reach  their  objective  point,  the  cells  change  their  form.  On  the  surface  appear  small 
humps  and  jagged  protuberances.  These  projections  later  elongate  and  become  compact 
branches  beset  with  small  knobs.  By  the  further  development  of  the  knobs  and  the  manifold 
division  oi  the  outgrowths,  the  later  protoplasmic  processes  or  dendrites  of  the  cells  arise. 
In  this  manner  the  nerve-cell,  or  rather  the  neurone,  is  formed  as  an  independent  individual. 
It  includes  the  cell-body  and  the  outgrowing  protoplasmic  processes  or  dendrites  and  sends  out 
the  delicate  netve-process  or  neurite,  which  in  its  later  development  becomes  the  nerve-fibre. 

DEVELOPMENT  OF  THE  CELLS  OF  THE  CEREBRO-SPINAL   GANGLIA   AND  THE 
SYMPATHETIC  GANGLIA. 

The  spinal   ganglia  are  developed  from  a  band  of   ectodermic  cells  located  where 
the  medullary  plate  passes  into  the  cuticle-layer.      In  the  stage  of  the  medullary  groove, 

this  ganglion-strand  occupies,  on  each  side,  the 
prominent  ridge  of   the   medullary  plate   and,  as 

spinal  ganglion  &=  the  medullary  tube  separates  from  the  overlying 

ectoderm,   unites   temporarily  with  the  strand  of 
the    other  side  to  form  a   common    medial  cord. 

FIG.    104. —  Development    of    the    ganglion-strand. 

Schematic.  In  consequence  of   the   formation  of   the   medul 


FIG.  103. — Further  development  of  the 
neuroblasts.  On  the  right,  two  neuroblasts 
exhibit  processes  bearing  growth-wedges. 


io6 


THE   FIBRE-TRACTS. 


lary  tube,  the  elements  of  the  ganglion-strand,  the  ganglioblasts,  are  displaced  laterally 
and  form,  on  each  side  of  the  medullary  tube,  segmentally  arranged  cell-groups.  The 
latter  are  the  anlagen  of  the  future  spinal  ganglia.  During  their  migration  along  the 
medullary  tube,  the  ganglioblasts  become  spindle-shaped.  This  form  becomes  subse- 
quently still  more  pronounced,  each  of  the  two  pointed  ends  gradually  growing  out 
into  a  nerve-fibre,  the  centrally  directed  one  growing  into  the  dorsal  portion  of  the 


Spinal  ganglion. 


Periph, 


G.  105. — Neuroblasts  and  ganglioblasts. 


cord  as  a  posterior  root-fibre,  and  the  other,  as  the  peripherally  directed  sensory  fibre, 
traversing  the  body  to  its  termination.  The  bipolarity  of  the  ganglion-cells  later  disappears, 
the  cells  becoming  unipolar  elements.  This  unipolarity  is  manifested  not  only  by  the  cells  of 
the  spinal  ganglia,  since  the  cells  of  the  corresponding  ganglia  of  the  cerebral  nerves  are  also 
unipolar  elements.  The  ganglion  acustici  alone  contains  permanently  bipolar  cells. 

The  sympathetic  ganglia  originate  from  the  cerebro-spinal  ganglia.  According 
to  the  younger  His,  this  development  is  accomplished  by  an  actual  migration  of  cellular 
elements  from  the  spinal  ganglia. 

THE   FORMED   ELEMENTS   OF  THE   NERVOUS  SYSTEM. 

The  formed  elements  of  the  nervous  system  are  the  support-cells  and  the  nerve-cells. 

A.    THE   SUPPORT-CELLS. 

These    include   the  ependyma   cells  and  the  neuroglia  cells.     The  former  constitute 
the  epithelial  lining,  the  ependyma,  of  the  central  canal  and  its  prolongations — the  fourth 
ventricle,  the  aquaeductus  cerebri,   the  third  ventricle  and  the 
lateral  ventricles. 

The  neuroglia  cells,  the  spider  cells  or  astrocytes,  are 
present  in  all  parts  of  the  gray  and  white  substance  and  form, 
by  means  of  their  numerous  processes,  the  framework  proper, 
the  astropilemma  or  spongiopilemma.  As  chief  forms,  we  dis- 
tinguish the  short-rayed  and  the  long-rayed  neuroglia  cells. 
They  all  possess  numerous  processes,  which,  however,  are 
seldom  uniformly  distributed  around  the  circumference  of  the 
cell-body,  but  usually  emerge  in  separate  close  tufts,  like  bun- 
dles of  rays.  The  processes  are  delicate,  mostly  of  the  same 

thickness,  from  beginning  to  end  of  uniform  width  and  end  free.  While  in  the  majority 
of  cells  they  proceed  in  all  directions,  there  are  also  astrocytes  in  which  the  processes 
exhibit  a  one-sided  development,  or  arise  from  the  two  poles  of  the  cell. 


FIG.    106. — Neuroglia    cells    from 
the  human  cerebral  cortex. 


THE   NERVE-CELLS.  107 

For  a  long  time  the  supporting  tissue  proper  of  the  nervous  system,  or  the  neu- 
roglia,  as  distinguished  from  connective  tissue,  blood-vessels  and  lymphatics,  was  regarded 
as  a  sort  of  ground-substance  in  which  the  nerve-cells  and  nerve-fibres  were  embedded. 
The  chief  r61e  therein  was  played  by  a  kind  of  cement-substance,  the  glia,  to  which  be- 
longed special  cells  and  fibrous  elements,  the  glia  cells  and  glia  fibres.  In  1811  Keuffel 
first  succeeded  in  demonstrating  a  definite  meshwork  in  cross-sections  of  the  spinal  cord, 
by  brushing  out  the  nervous  sub'stance,  and  believed  that  this  reticulum  represented  noth- 
ing more  than  the  prolongations  of  the  pia  mater.  Arnold  and  Virchow  termed  the  neu- 
roglia  a  granular  ground-substance,  but,  as  early  as  1853,  Virchow  demonstrated  round 
or  fusiform  cells  within  this  ground-mass  and  regarded  the  tissue  as  of  nervous  character, 
believing  that  the  nerve-cells  were  developed  from  the  neuroglia.  Bidder  went  somewhat 
further  and  spoke  of  fibrillae  and  stellate  cells  with  processes.  In  1863  Kolliker  pointed 
out  that  the  supporting  tissue  of  the  nervous  system  consisted  of  nothing  else  than  a 
complex  of  anastomosing  stellate  cells,  which  by  their  union  formed  a  reticulum  for  the 
nervous  elements.  He  still  assumed,  however,  an  anastomosis  between  the  processes  of 
the  cells.  It  remained  for  Deiters,  by  means  of  isolation,  to  represent  the  neuroglia  cells 
in  their  correct  form.  The  greatest  service,  however,  was  rendered  by  Golgi.  Through 
his  investigations,  it  became  clear  that  the  neuroglia  is  not  a  special  issue,  but  that  it  is 
represented  by  certain  elements — the  neuroglia  cells,  spider  cells  or  astrocytes. 

B.    THE   NERVE-CELLS. 

The  first  accurate  description  of  the  nerve-cell  was  given  by  Remak  in  1838. 
Thirteen  years  later,  R.  Wagner  discovered  in  the  nerve-cells  of  the  electrical  lobes  in 
the  brain  of  torpedo,  that  of  the  processes  passing  out  from  the  cells  only  a  single  one 
is  connected  with  a  nerve-fibre.  In  1854  Remak  communicated  similar  results  in  his 
studies  on  the  nerve-cells  of  the  gray  ventral  columns  of  the  spinal  cord  of  the  ox. 
These  observations  of  Wagner  and  of  Remak  were  confirmed,  in  1865,  by  Deiters'  in- 
vestigations on  the  human  brain  and  spinal  cord.  Deiters  found  that  among  the  many 
processes  passing  from  a  nerve-cell  always  one  ran  unbranched,  while  the  others  under- 
went repeated  division.  The  unbranched  process  he  named  the  nerve-process  or  axis- 
cylinder  process,  and  the  branched  ones  the  protoplasmic  processes.  In  his  investigations 
Deiters  employed  the  method  of  isolation,  this  teasing  method  being  subsequently  long 
used  to  demonstrate  the  nerve-cells.  It  is  evident,  however,  that  with  such  technique, 
by  which  the  cells  were  torn  from  all  their  relations,  other  investigators  could  achieve 
little  more  than  Deiters  had  already  done,  and  that  the  most  diverse  statements  c.oncern- 
ing  the  conception  of  the  relations  of  adjacent  elements  to  each  other  were  inevitable. 
Thus,  a  direct  union  of  neighboring  cells  with  each  other  was  accepted  by  many  investi- 
gators as  an  undoubted  fact.  Sometimes  it  concerned  broad  connecting  bridges  or  anas- 
tomoses, sometimes  the  passage  of  delicate  end-fibres  into  each  other.  According  to 
other  investigators,  all  nerve-cells  possessed  more  than  a  single  typical  nerve-process. 
Gerlach's  work  merits  the  greatest  consideration.  Gerlach  succeeded  in  demonstrating 
an  exceptionally  rich  felt-work  of  the  most  delicate  nerve-fibres  in  all  parts  of  the  gray 
substance.  He  extended  the  observations  of  Deiters,  who  had  seen  the  protoplasmic 
processes  repeatedly  branch  and  also  the  most  delicate  of  these  ramifications  still  further 
subdivide,  in  that  he  held,  that  the  finest  ramifications  of  the  protoplasmic  processes 


io8  THE   FIBRE-TRACTS. 

eventually  formed  a  delicate  "nerve-fibre  reticulum."  This  Gerlach  regarded  as  the  most 
important  constituent  of  the  gray  substance.  According  to  Gerlach,  the  divisions  of  the 
delicate  protoplasmic  processes  observed  by  Deiters  were  only  the  beginning  of  the  ner- 
vous reticulum.  Gerlach,  however,  went  still  further.  He  assumed  that  from  this  net- 
work of  nerve-fibres,  by  the  gradual  confluence  of  the'  minute  branches,  broader  nerve- 
fibres  were  again  formed,  which  emerged  from  the  gray  substance.  Accordingly,  the 
nerve-fibres  had  a  two-fold  origin,  on  the  one  hand,  directly  from  the  cells  as  nerve- 
processes  or  axis-cylinder  processes,  and,  on  the  other,  indirectly  from  the  cells  through 
the  medium  of  the  reticulum  of  nerve-fibres  resulting  from  the  branching  of  the  proto- 
plasmic processes.  Gerlach  supposed,  therefore,  that  the  end-twigs  of  the  sensory  fibres 
entered  the  delicate  network,  which,  on  the  other  side,  received  the  branched  protoplas- 
mic processes  of  the  motor  nerve-cells.  Gerlach' s  fibre-reticulum  can  be  best  pictured  by 
comparing  it  with  the  capillary  network  of  the  blood-vessels ;  the  sensory  fibre  is  the 
artery,  which  is  resolved  into  the  capillary  network ;  the  protoplasmic  processes  of  the 
cells  form  the  beginnings  of  the  venous  reticulum,  from  which  proceeds  the  nerve-process 
representing  the  vein. 

This  nerve-fibre  reticulum  of  Gerlach  enjoyed  for  a  long  time  general  acceptance. 
With  the  improvements  in  the  methods  of  investigation,  however,  a  powerful  revulsion 
took  place.  In  this,  the  chief  role  was  played  by  the  silver-method  of  Golgi.  This 
investigator  made  the  important  discovery,  that  the  nerve-processes  of  the  cells,  regarded 
as  unbranched,  may  give  off  delicate  collateral  branches.  Moreover,  that  there  are  many 
cells  in  the  brain  and  spinal  cord  whose  nerve-processes  are  riot  continued  as  medullated 
nerve-fibres,  as  in  the  case  of  the  other  cells  and  in  conformity  with  the  general  law 
announced  by  Deiters,  but  resolve  into  their  ultimate  end-twigs  immediately  after  emerg- 
ing from  the  cells  or  after  a  short  course. 

Golgi  divided,  therefore,  the  nerve-cells  of  the  brain  and  the  spinal-cord  into  two 
classes  :  Type  I,  cells  with  long  nerve-processes,  which  latter  are  directly  prolonged  into 
nerve-fibres  ;  Type  II,  cells  with  short  nerve-processes,  which  after  a  short  course,  almost 
immediately  after  exit  from  the  cells,  break  up  into  their  terminal  branches. 

Later,  the  two  varieties  were  described  as  the  types  of  Deiters  and  of  Golgi.  Also 
functionally  these  two  cell-types  differ.  Golgi  regarded  the  Deiters  cells  as  motor  and 
the  others  as  sensory  elements.  He  interpreted  the  protoplasmic  processes  as  merely  the 
nutritive  organ  of  the  nerve-cell  and  questioned  their  nervous  significance.  Of  most 
importance,  however,  is  the  hypothesis  advanced  by  Golgi  and  his  pupils  concerning  the 
internal  connection  of  the  central  nervous  apparatus.  Golgi  denied  anastomoses  of  the 
protoplasmic  processes  with  one  another  and,  consequently,  a  connection  between  the 
cells  in  the  sense  of  Gerlach,  although  suggesting  a  view  somewhat  similar.  He  cham- 
pioned the  existence  of  a  "general  nervous  network,"  which,  on  the  one  hand,  arises  from 
the  delicate  collateral  branches  of  the  long  nerve-processes  and  from  the  terminal  subdi- 
visions of  the  nerve-processes  of  cells  assumed  by  him  to  be  sensory  elements,  and,  on 
the  other  hand,  receives  additional  constituents,  such  as  the  end-twigs  of  the  nerve-fibres 
which  enter  the  gray  substance.  This  network  he  believes  to  exist  throughout  the  entire 
gray  substance  of  the  spinal  cord  and  of  the  brain. 

Opposed  to  this  ' '  nervous  network' '  are  the  important  considerations  of  His  and 
of  Forel.  As  early  as  1883,  based  upon  embryological  investigations,  His  had  shown 


THE   NERVE-CELLS. 


109 


the  independence  of  the  nerve-cells  from  one  another ;  while  in  1887,  chiefly  upon 
pathological  experiences  with  Gudden's  atrophy-method,  Forel  opposed  the  acceptance  of 
a  general  network.  What  he,  for  the  first  time,  especially  emphasized,  was  the  principle 
of  contact  instead  of  continuous  reticular  connection.  There  was  still  wanting,  however, 
the  histological  proof,  and  this  proof  was  supplied  by  the  Spanish  savant,  Ram6n  y  Cajal. 
By  means  of  his  investigations,  it  was  established  that  every  nerve-cell,  with  its  emerg- 
ing nerve-fibres,  represented  an  histological  as  well  as  a  neurological  entity— a  neurone, 
— and  that  the  entire  nervous  system  is  built  up  of  such  nervous  units. 

Closer  examination  of  such  a  nervous  unit  or  neurone  (Fig.  107),  shows  that  two 
kinds  of  processes  leave  the  cell-body  :  (a)  the  branching  protoplasmic  processes  or  the 
dendrites  and  (6)  the  axis-cylinder  process,  also  called 
the  nerve-process,  axone  or  neurite.  The  nerve-process 
is  distinguished  by  its  uniform  diameter  and  smooth, 
regular  structure.  During  its  course  it  gives  off  many 
secondary  twigs,  the  collaterals  or  paraxones,  and  ends 
by  forming  a  terminal  arborization  or  telodendrion.  All 
these  parts — the  cell  with  its  dendrites  and  the  axone 
with  its  telodendrion — constitute  collectively  a  nervous 
unit  or  a  neurone. 

Concerning  the  function  of  the  individual  parts  of 
the  neurone,  the  cell-body  with  its  dendrites  forms  the 
perceptive  and  impulse-giving  element,  while  the  nerve- 
process  with  its  collaterals  and  the  end-arborization,  is 
the  organ  of  transmission,  carrying  the  impulse  from  the 
nerve-cell  to  other  elements.  The  protoplasmic  proc- 
esses or  dendrites,  therefore,  conduct  cellulipetally , 
receiving  the  impulse  and  carrying  the  same  to  their 
own  cell-body  ;  the  nerve-process  or  neurite  conduct 
cellulifugally ,  receiving  the  nervous  stream  from  its 
own  cell-body  and  conducting  it  to  other  cells. 

The  manner  in  which  the  transference  from  one 
neurone  to  the  other  occurs  is  not  definitely  known. 
According  to  certain  investigators,  the  chaining  together 

of  the  nervous  units  is  effected  by  the  nerve-process  of  one  cell,  split  into  the  delicate 
fibres  of  its  end-arborization,  being  closely  applied  to  or  overlying  the  dendrites  and 
cell-body  of  another  cell,  whereby  the  transference  of  the  impulse  is  accomplished. 
The  opponents  of  the  theory  of  mere  contact  hold  that  there  exists  not  only  a  simple 
application,  but  also  a  continuous  connection  between  the  nervous  substance  of  the  nerve- 
processes  and  of  the  protoplasmic  parts  in  the  form  of  an  extremely  delicate  nervous 
network.  Even  if  definite  proof  were  supplied  as  to  the  connection  of  the  individual 
neurones  with  one  another  by  means  of  such  a  network,  it  remains  none  the  less  certain, 
that  the  nerve-cells  with  their  processes  are  the  essential  elements  for  the  entire  nervous 
activity  and  that  they  must  be  regarded  as  the  elements  of  the  nervous  system,  which 
anatomically,  trophically  and  as  regards  specific  function,  enjoy  a  certain  degree  of  indepen- 
dence. We  are,  therefore,  justified  in  designating  them  as  nervous  units  or  neurones. 


FIG.    107. — Sche 


tic   representation  of   a 
:urone. 


no 


THE   FIBRE-TRACTS. 


The  nerve-cells  are  found  chiefly  in  the  central  nervous  system  ;  further,  in  the 
ganglia,  the  sense-organs  and  in  the  course  of  the  cerebro-spinal  and  the  sympathetic 
nerves.  They  are  of  variable  size  (4  to  135/0  and  of  manifold  shape.  The  chief  charac- 
teristic of  every  nerve-cell  consists  in  its  always  possessing  processes.  Nerve- cells  with- 
out processes,  the  so-called  apolar  cells,  are  never  found  in  the  nervous  system  of  the 
adult.  Such  cells  are  either  immature  forms  and  found  only  during  the  earliest  period 
of  embryonic  development,  as  the  germ-cells  of  His,  or  they  are  artificial  products,  arising 
from  the  tearing  off  of  the  processes  during  isolation. 

According  to  the  number  of  the  processes,  unipolar,  bipolar  and  multipolar  cells 
are  distinguished. 


FIG.  108. — Nerve-cells  of  different  types,    a,  unipolar  cells;    b,  bipolar  cells;    c,  pyramidal  cell;    d,  Purkinje  cell. 

Unipolar  Cells.  These  are  numerous  during  embryonic  development,  as  the 
neuroblasts  ;  much  less  frequently  they  are  encountered  in  the  nervous  system  of  the 
adult,  as  in  the  retina  and  in  the  mesencephalon  on  each  side  of  the  aquaeductus  cerebri 
as  the  cells  of  origin  of  the  upper  motor  root  of  the  nervus  trigeminus.  The  nerve- 
cells  of  the  cerebro-spinal  ganglia  are  apparently  unipolar,  with  the  exception  of  the  cells 
of  the  ganglion  spirale  and  of  the  ganglion  Scarpae  ;  in  their  embryonic  condition,  how- 
ever, these  elements  are  bipolar,  only  later  becoming  unipolar,  when  their  nerve-processes 
divide,  at  a  certain  distance  from  the  cell-bodies,  into  a  central  and  a  peripheral  branch. 

Bipolar  Cells.  These  elements  are  found  almost  exclusively  in  the  peripheral 
sensory  nervous  system,  as  in  the  epithelium  of  the  olfactory  mucous  membrane,  in  the 
retina  and  in  the  spinal  and  vestibular  ganglia. 

Multipolar  Cells.  These  are  the  most  numerously  represented  and  the  most  impor- 
tant elements  of  the  nervous  centres.  Connected  with  these  are  two  kinds  of  processes — 
the  nerve-process,  axis-cylinder  or  neurite,  and  the  protoplasmic  processes  or  dendrites. 


THE   NERVE-CELLS. 


Dendrites 


FIG.  109. — Nerve-cell    from  spinal   cord   of  a  new 
born  cat. 


The  nerve-process  or  neiirite  is  usually  single,  although  cells  with  several  nerve- 
processes  occur  within  the  central  cortex,  as  the  cells  of  Cajal.  To  this  category  belong 
also  the  multipolar  cells  of  the  sympatheticus  described  by  different  authors.  The  neurite 
leaves  the  cell  by  means  of  a  small  conical  elevation,  the  implantation  cone ;  the  origin 
may  be  either  from  the  cell,  or,  as  is  very  often  the  case,  from  one  of  the  protoplasmic 
processes,  near  or  at  some  distance  from  the  cell-body.  Its  smooth  regular  quality  and 
uniform  diameter  throughout  its  entire  course  are  characteristic  of  the  nerve-process. 

The  protoplasmic  processes  or  the  dendrites  are  broad  and  dense  at  their  origin  from 
the  cell-body,  become  gradually  thinner  and  repeatedly  undergo  antler-like  division  to 
form  often  an  arborization  of  extraordinary  richness,  the  finest  twigs  of  which  end  free. 

Their  irregular  course  and  knobbed  condition 
are  characteristic  of  the  dendrites,  which  are 
often  beset  with  numerous  knots,  thorns  or  spines. 
According  to  the  behavior  of  their  nerve- 
processes,  nerve-cells  are  grouped  into  two  classes 

i.  Cells  with' long 
nerve  -processes,  Deiters' 
cell-type  (Fig.  109),  in 
which  the  neurite  is  ex- 
tremely long  and  becomes 
the  axis-cylinder  of  a  cen- 
tral or  'peripheral  nerve- 
fibre. 

2.  Cells  with  short  nerve-processes,  Golgi's  cell-type  (Fig. 
no),  in  which  the  short  neurite  does  not  become  a  nerve- 
fibre,  but,  close  to  the  cell,  undergoes  repeated  division  and 

resolves  into  its  end-arborization.  These  elements  are  conveniently  designated  as 
Golgi  cells,  or  cells  of  Golgi's  Type  II,  as  distinguished  from  the  cells  of  Golgi's 
Type  I,  or  the  cells  with  long  neurites. 

According  to  the  behavior   of  the  protoplasmic   processes,  are  distinguished: 

a.  Stellate   cells,  the  dendrites  of  which  arise  separately  from  the  entire  circumfer- 
ence of  the  cell-body  and  extend  in   all    directions,    as  the  motor  anterior   horn-cells  and 
the  tract-cells  of  the  spinal    cord. 

b.  Cells  with  a  chief  dendrite,  in  which  a  robust  protoplasmic  process  arises,  along 
with  other  dendrites,  gives  off  lateral  branches  and  ends  arborized,  as  the  pyramidal  cells 
of  the  cerebral  cortex  and  the  mitral  cells  of  the  bulbus  olfactorius. 

c.  Cells  with  polar  dendrites,  in  which  the  cell-body  is    mostly  fusiform  and  gives 
off  from  opposite  sides  a  basal  and   an  apical   dendrite.     The  basal  dendrite  forms  a  tuft 
resembling  the  roots  of  a  tree,  while  the  apical  dendrite  springs  from  a  chief  protoplasmic 
process,  which  eventually  likewise   resolves   into   numerous  branches.     The  nerve-process 
springs  from  a  basal  dendrite,  as  in  the  pyramidal  cells  of  the  hippocampus. 

d.  Cells  with   monopolar    dendrites,    in    which  usually   several   chief   dendrites  arise 
from  one  pole   of    the    cell-body  and  soon  break-up,  after   repeated  division,  into  a  wide 
arborization.     The  nerve-process  arises  from  the  other  pole,  as    in    the    Purkinje    cells  of 
the  cerebellum,  or  in  the  granule  cells  in  the  gyrus  dentatus. 


FIG.  no.— Nerve-cell 
with  short  axone  or  nerve- 
process;  cerebral  cortex. 


H2  THE   FIBRE-TRACTS. 

"With  regard  to  their  intimate  structure,  the  nerve-cells  may  be  divided  into  two 
chief  groups,  in  accordance  with  the  behavior  of  their  protoplasm  towards  the  basic 
anilin  dyes.  Following  Nissl,  we  distinguish  somatochromic  and  karyochromic  cells ;  in 
the  former  both  nucleus  and  protoplasm  stain,  in  the  latter  only  the  nucleus.  After 
staining  with  basic  anilin  colors,  such  as  methylene  blue  or  thionin,  the  protoplasm  of  the 
somatochromic  cells  exhibits  a  part  taking  the  dye,  the  chromophilic  substance, 
and  a  part  remaining  uncolored,  the  chromophobic  substance.  The  stainable  part  appears 
as  a  multitude  of  deeply  colored  bodies  having  the  form  of  spherical  granules,  threads, 
flakes,  spindles  or  jagged  particles,  which  also  extend  into  the  dendrites,  but  do  not 
invade  the  axis-cylinder  process.  These  are  known  as  Nissl  bodies  or  granules.  On 


PIG.  in. — Structural  details  of  nerve-cells,  a,  pyramidal  cell  from  adult  human  motor  cortex,  showing  neurofibrillar 
network;  6,  pyramidal  cell  from  adult  human  visual  cortex;  c,  anterior  horn-cell 'from  human  spinal  cord  showing  Nissl 
bodies;  d,  Golgi-Holmgren  canals  in  pyramidal  cell  of  rabbit;  e,  ending  of  nerve-fibres  on  nerve-cells;  ,/,  pericellular  or 
Golgi  network,  (a.  6,  d,  e,  and  /  after  Cajal;  c,  after  Schmaus.) 

account  of  the  mottled  appearance  of  the  cell-body  conferred  by  the  staining  substance, 
von  Lenhosse'k  calls  the  latter  tigroid.  The  arrangement  of  the  chromophilic  substance 
is  variable,  the  granules  sometimes  being  irregularly  scattered,  at  other  times  disposed 
in  concentric  layers,  or,  as  in  the  case  of  fusiform  cells,  grouped  as  a  sort  of  cap  at 
each  pole  of  the  cell-nucleus.  A  conical  mass  of  the  chromophilic  substance  is  usually 
found  at  the  points  of  division  of  the  dendrite-stems.  Concerning  the  chromophilic  part 
of  the  protoplasm,  the  nerve-fibrillae  or  neurofibrillae  deserve  first  notice.  These  occur 
within  the  cell-body,  as  well  as  within  the  processes,  and  constitute  a  more  or  less  ex- 
tensive reticulum,  the  fibrillar  network.  In  Fig.  in,  a  and  b,  such  intracellular 
networks  are  represented.  The  figures  e  and  f  show,  further,  how  nerve-fibres,  after 
resolving  into  delicate  arborizations,  end  at  the  cells  and  how  the  finest  fibres  form  a 
delicate  network  over  the  surface  of  the  cell-body  and  the  dendrites,  this  constituting 
the  external  network  of  Golgi. 


THE   NERVE-CELLS.  113 

The  chromophilic  and  the  chromophobic  substance  differ  also  in  their  functional 
relations.  The  chromophilic  substance  is  wanting  in  the  protoplasm  of  a  large  number 
of  nerve-cells  and,  therefore,  does  not  represent  a  vital  element  of  the  nerve-cell.  It 
accumulates  during  the  resting  condition,  is  sometimes  notably  reduced  during  the  period 
of  activity  and  disappears  in  lesions  'of  the  neurone,  to  reappear  in  generous  quantity, 
however,  after  temporary  injury  and  recovery  of  the  cell.  These  characteristics  seem  to 
prove,  that  the  chromophilic  substance  exercises  a  nutritive  rather  than  a  nervous 
function.  The  chromophobic  substance  seems  to  represent  the  element  possessing  the 
function  of  conducting  the  nervous  stream,  the  fibrillae  and  the  perifibrillar  substance 
probably  sharing  in  this  conduction. 

In  addition  to  the  Nissl  bodies,  the  protoplasm  of  many  cells  contains  pigment 
granules,  whi£h  are  usually  disposed  in  groups  of  variable  size.  The  pigment  is  com- 
monly not  uniformly  distributed  within  the  cell,  but  arranged  at  the  base  of  one  of  the 
dendrites.  It  is  wanting  during  early  life  and  increases  with  age.  Marinesco  regards 
the  pigment  granules  as  regression-  and  age-products  of  the  nerve-cells.  Claim- 
ing mention  are,  further,  the  fine  channels,  the  canals  of  Holmgren,  which  lie 
within  the  interior  of  the  cell  and  communicate  -with  the  lymph-canals  situated  outside 
the  nerve-cell. 

The  nucleus  of  the  nerve-cells  appears  as  a  clear  spherical  vesicle,  possesses  a  distinct 
nuclear  membrane  and  lies  most  frequently  in  the  middle  of  the  cell.  Within  the  nucleus  are 
found  one  or  several  deeply  staining  nucleoli,  which  often  contain  still  smaller  bodies, 
the  nucleololi.  The  remaining  interior  of  the  nucleus  is  traversed  by  a  meagre  sup- 
porting substance,  the  linin  framework,  on  which  rests  the  chromatin,  as  well  as  against 
the  nuclear  membrane. 

Concerning  the  relations  of  the  cells  to  the  surrounding  tissue,  it  should  be  noted, 
that  the  cells  are  enclosed  within  cavities,  the  pericellular  spaces,  which  communicate  with 
the  perivascular  lymph-channels  of  the  central  nervous  system. 

The  envelopes  of  the  nerve-cells  are,  according  to  Cajal,  of  two  kinds,  (a)  The 
cell-membrane  proper,  the  membrana  fundamental,  which  invests  every  cell  of  the  gray 
substance  as  an  extremely  delicate,  homogeneous,  elastic  cuticle,  and  (^)  a  connective 
tissue  envelope,  a  delicate  nucleated  membrane,  which  surrounds  all  the  peripheral  cells — 
ganglion  cells  and  the  cells  of  the  sympathetic — with  the  exception  of  the  cells  of  the 
retina  and  of  the  olfactory  mucous  membrane. 

The  ependyma  and  the  neuroglia  cells  are  sustentacular  elements  and  together  form 
the  supporting  framework  of  the  nervous  system. 

The  nerve-cells  are  usually  closely  placed  in  larger  or  smaller  groups  and  consti- 
tute the  essential  components  of  the  gray  masses  of  the  nervous  system ;  exceptionally 
they  occur  singly,  scattered  within  the  white  substance. 

The  nerve-fibres  are  the  axis-cylinder  processes  or  nerve-processes  of  the  nerve- 
cells,  and,  while  everywhere  encountered  within  the  gray  masses,  constitute  the  chief 
components  of  the  white  substance  of  the  nervous  system.  They  serve  to  establish  rela- 
tions of  the  nerve-cells  with  one  another,  whether  the  relations  be  between  neighboring 
or  widely  separated  cells  of  one  and  the  same  region  of  gray  substance,  as  between 
the  various  regions  of  the  cerebral  cortex;  whether  the  relations  be  between  the  cells 
of  a  certain  region  and  those  of  one  far  remote,  as  between  the  central  cortex  and  the 

8 


u4  THE   FIBRE-TRACTS. 

deeper  placed  gray  masses  (thalamus,  pons,  medulla,  oblongata  and  spinal  cord);  or 
whether  the  relations  be  between  the  central  and  the  peripheral  nervous  system. 

The  nerve-cells  are,  therefore,  the  specific  function-bearing  elements.  They  are  the 
force-sources  or  the  transposition-apparatus  of  the  various  forms  of  nervous  activity  and, 
at  the  same  time,  also  the  nutritive  organs,  the  trophic  or  nutritive  centres,  of  the  nerve- 
fibres  which  pass  from  the  cells.  A  nerve-fibre  separated  from  its  nutritive  centre  loses 
its  function  and  no  longer  conducts.  A  nerve- cell  with  its  protoplasmic  processes,  or 
dendrites,  and  its  nerve-process,  or  neurite,  constitutes  a  nervous  unit,  or  a  neurone. 
The  protoplasmic  process  conducts  cellulipetally ;  the  nerve-process  conducts  cellulifugally 
and  through  it,  by  means  of  its  end-arborization,  as  well  as  of  its  collaterals,  occurs  the 
transference  of  the  impulse  from  one  neurone  to  the  other. 

Cells  of  the  same  function  usually  lie  together,  closely  packed,  and  constitute  a  region, 
a  centre,  a  ganglion  or  a  nucleus.  In  like  manner  fibres  of  the  same  function  usually  lie 
together,  closely  placed,  and  form  a  path  of  conduction  or  a  fibre-system. 


MICROSCOPIC  STRUCTURE  OF  THE  CEREBRAL  CORTEX. 

I.   CORTEX  OF  THE  PALLIUM. 

Based  on  the  arrangement  of  the  nerve-cells,  the  following  six  strata  or  layers  are 
distinguished. 

i.  The  molecular  layer.  This  constitutes  the  most  superficial  stratum  and  is  a 
dense  feltwork,  composed  principally  of  fibres  running  mostly  parallel  to  the  surface  ;  hence, 


/.  Molecular  laye 


II.  Outer  grannie 
layer 


III.  Layer  of  small 
Pyramidal  cells 


V.  Inner  granule         fc\ 
layer  [  /. 


V.  Layer  of  large 
Pyramidal  ctlls 


VI.  Layer  of  poly- 
morphic cells 


White  substance 


FIG.  112. — Schematic  representation  of  the  structure  of  the  cerebral  cortex. 


STRUCTURE  OF  CEREBRAL  CORTEX.  115 

it  is  also  designated  as  the  layer  of  tangential  fibres  or  the  tangential  fibre-layer.  In 
addition  to  numerous  neuroglia  cells,  this  layer  contains  the  terminal  ramifications  of 
the  dendrites  of  the  more  deeply  lying  pyramidal  cells  and  the  end-arborizations  of  the 
nerve-fibres  coming  from  the  white  substance  and  ending  in  the  cortex.  Further,  it  con- 
tains certain  cells,  including  medium  sized  polygonal  elements,  with  from  four  to  six 
dendrites  and  a  nerve-process  arborizing  within  the  molecular  layer,  and  fusiform  or  tri- 
angular cells,  with  few  more  or  less  horizontally  coursing  dendrites  and  two  or  several 
nerve-processes,  that  also  run  horizontally  and  end  within  the  molecular  layer.  The  ele- 
ments with  several  neurites  encountered  within  the  tangential  fibre-layer,  are  known 
as  Cajal  cells. 

2.  The  outer  granule  layer,  a  stratum  of  small  pyramidal  cells. 

3.  The  layer  of  small  arsd  medium  sized  pyramidal  cells. 

4.  The  inner  granule  layer,  a  stratum  of  small  pyramidal  cells. 

5.  The  layer  of  large  pyramidal  cells.     The  cell-body  of  the  pyramidal  cells  is 
pyramidal  in  form,  the  base  presenting  towards  the  white  substance  and  the  apex  directed 
towards  the  molecular  layer.     The  apex  is  prolonged  into  a  robust  protoplasmic  process, 
the  primordial  branch,  which   gives    off   lateral    twigs   at   right   angles,  runs   toward   the 
molecular  layer  and  there  ends  after  repeated  division.     The  basal  dendrites  pass  off  from 
the   base  of  the  cell-body,  radiating  laterally,  or  towards  the  white   matter.     The  nerve- 
process  springs  from  the  base  of  the  cell,  or  from  a  basal  dendrite  close  to  the  cell-body, 
and   runs   towards   the   white  substance ;    during   its   course   through  the  gray  substance, 
the  nerve-process   gives   off   fine   collaterals,  that   run   horizontally  or  obliquely    and   end 
after  a  number  of  branchings. 

6.  The  layer   of  polymorphic  cells.      Here   are  found   cells,  ovoid,    fusiform, 
triangular    or   polygonal    in    form,    which    often    exhibit   a   robust   protoplasmic    process, 
directed  towards  the  molecular  layer.       Each   cell  sends   off   a   nerve-process   that  passes 
to  the   white  substance,  after  giving  off  a  number  of   collaterals.      Additional    cells,   with 
short  nerve-processes  or  of  Golgi's  II  type,  are  encountered,   not  only  in  this  layer,   but 
also  within  the  strata  of  small  and  large  pyramidal  cells.       Finally,   the  so-called  cells  of 
Martinotti  occur   as   fusiform   or   triangular    elements,  whose   distinguishing   characteristic 
consists  therein,  that  the  nerve-process  traverses  the  layer  of  the  pyramidal  cells  to  reach 
the  molecular  stratum,  where  it  ends. 

Regarding  the  disposition  of  the  nerve-fibres,  thicker  or  thinner  parallel 
bundles  of  fibres  enter  the  cortex  from  the  white  substance,  proceed  towards  the  periph- 
ery, and,  gradually  diminishing  in  thickness,  towards  the  layer  of  the  small  pyramidal 
cells  resolve  into  their  component  fibres.  These  bundles  are  known  as  the  medullary 
rays  or  radii  and  consist  of  the  nerve-processes  of  the  pyramidal  and  of  the  polymorphic 
cells,  which  are  passing  from  the  cortex,  and  of  the  nerve-fibres,  which  enter  from  the 
white  substance  and  end  within  the  cortex;  these  last,  are  also  called  the  terminal  fibres. 
Between  the  individual  medullary  rays  are  narrow  interspaces  containing  delicate  horizon- 
tally coursing  fibres,  which  form  the  interradial  feltwork.  The  latter  are  somewhat 
denser  where  the  medullary  radii  break  up  into  their  individual  fibres  and  thereby  pro- 
duce the  stripe  of  Baillarger.  The  fibres  of  the  interradial  feltwork  are  the  collaterals 
of  the  nerve-processes  of  the  pyramidal  cells.  Towards  the  periphery,  beyond  the  inter- 
radial feltwork  where  the  radii  resolve  into  their  component  fibres,  lies  the  supraradial 


n6 


THE   FIBRE-TRACTS. 


feltwork,   which  marks  the  ending  of  the  terminal  fibres   and,  farther   outward,  joins  the 
layer  of  tangential  fibres. 

The  cerebral  cortex  does  not,  however,  present  the  same  structure  in  all  regions. 
Local  variations  occur,  in  relation  to  the  arrangement  of  the  several  cell-layers,  as  well 
as  in  regard  to  the  behavior  of  the  fibre-layers.  There  exists  a  cyto-  and  a  myelo- 


liiu 


IP 
IS™ 

a ft li,ifl> ' 

SfBK&f 


PIG.  113. — Structure   of  the   cerebral  cortex  in  different  regions.     A,  cortex   of  precentral   convolution;  B,  postcen- 
(convolution;  C,  superior  temporal  convolution  (auditory  cortex);  D,  surrounding  the  calcarine  fissure   (visual  cortex). 

linl  1 


tral  convoluti 
(CajaJ.) 

architectonic  differentiation,  the  recent  investigations  of  Brodmann  and  of  Vogt  having 
shown  that  the  entire  cerebral  cortex  may  be  subdivided  into  numerous  histologically 
different  cortical  fields.  While  it  is  impracticable  here  to  discuss  in  detail  the  differ- 
ences, Fig.  113  presents  these  relations,  so  far  as  the  make-up  of  the  cell-layers  is  con- 
cerned, in  the  precentral  and  postcentral  convolutions  and  in  the  auditory  and  visual 
cortical  areas,  according  to  the  earlier  studies  of  Ramon  y  Cajal.  The  preponderance  of 


CORTICAL   AREAS.  117 

the  large  and  giant  pyramidal  cells  in  the  precentral  convolution  is  to  be  noted  in  contrast 
to  the  peculiar   structure   of   the  visual  cortex,  in   which  the   original  six-layered   type  is 


PIG.  114. — Lateral  surface  of  hemisphere,  with  cytoarchitectonic  cortical  areas.     (Brodmann.) 


FIG.  115.— Mesial  surface  of  hemisphere,  with  cytoarchitectonic  cortical  areas.     (Brodmann.) 

transformed  into  one  of  nine  layers,  by  the  introduction  of   special  layers  of  stellate  cells 
(Fig.    113,    04  and  5). 


n8 


THE  FIBRE-TRACTS. 


Layer  of  mitral  cells 


II.   RHINENCEPHALON. 

Microscopic  structure  of  the  bulbus  olfactorius,  the  gyrus  fornicatus,  the  hippocampus 
and  the  gyrus  dentatus. 

BULBUS   OLFACTORIUS. 

The  bulbus  olfactorius  exhibits  the  following  layers  : 

i.  The  layer  of  the  superficial  nerve-fibres.  This,  the  fibre-layer,  is  formed 
by  the  nerve-fibres  coming  from  the  olfactory  epithelium  (Fig.  116).  Within  the 
epithelium  of  the  olfactory  mucous  membrane,  the  bipolar  nerve^cells  lie  among  the  sus- 
tentacular  cells.  They  are  elongated  narrow  fusiform  or  irregular  elements,  with  a  thick 

peripherally  directed  process, 

Fibres  ending  m  bnllns        Grannie  cells         Golgi  cells  that  ends  within  the  epithelium, 

and  a  delicate  centrally  di- 
rected process,  beset  with 
varicosities,  that  traverses  the 
tunica  propria  undivided. 
United  into  small  bundles,  the 
fila  olfactoria,  the  central  fibres 
pass  through  the  apertures 
of  the  lamina  cribrosa,  enter 
the  bulbus  olfactorius  and 
there  form  a  close  feltwork  of 
crossing  fibres,  the  fibre-layer, 
2.  The  glomerular  lay- 
er. Joining  the  stratum  of 
nerve-fibres,  the  layer  of  glom- 
eruli  olfactorii  follows.  Here 
the  end-arborizations  of  the 
fibres  from  the  fibre-layer  meet 
those  of  the  dendrites  of  certain 
cells,  namely,  the  brush  and 

mitral  cells.  In  consequence  of  the  close  intermingling  of  these  delicate  terminal  twigs,  small 
round  or  ovoid  structures,  the  glomeruli  olfactorii,  are  formed.  The  olfactory  fibres  com- 
posing the  fibre-layer  divide  often  into  two,  or  indeed  into  three,  twigs,  which  enter  the 
glomeruli ;  in  this  manner,  such  ramifications  may  penetrate  into  two  different  glomeruli. 

3.  The  molecular  layer.    This  layer,  also  known  as  \.\\&  stratum  gelatinosum  of  Clarke, 
forms   a  stratum  comparable  to  the  layer  of  small  pyramidal  cells  of   the  cerebral  cortex. 
Within  it,  along  with  traversing  and  branching  fibres,  are  found  large  and  small  brush-cells. 

4.  The  layer  of  mitral  cells.     When  compared  with  the  cerebral  cortex,  it  rep- 
resents the  layer  of  the  large  pyramidal   cells.     The  component  mitral  cells  are  of  quite 
characteristic  form.       The  cell-body  is  large,   exhibits    the  form  of   a  triangle  or  a  mitre,  • 
and   resembles   that   of   the    Purkinje   cells   of   the   cerebellar   cortex.       The   protoplasmic 
processes    are    of   two   kinds,   the   ordinary  dendritic    stems   and    the    so-called    olfactory 
brushes,  the  penicilli  olfactorii.     The  former  pass  obliquely  from  the  cells,  then  run  more 


FIG.  116. — Olfactory   mucous   membrane  and   bulbus   olfactorius.     Schematic. 


GYRUS  FORNICATUS.  119 

horizontally,  branch  once,  and  end,  usually  after  a  long  course,  free,  forming  a  feltwork 
within  the  deepest  part  of  the  molecular  layer.  The  olfactory  brushes  traverse  tha 
molecular  layer  and  assist  in  forming  the  glomeruli  with  their  elaborate  varicose  end- 
arborizations.  The  nerve-processes  of  the  mitral  cells  extend  towards  the  granule-layer, 
bend  sagittally  at  various  levels  and  continue  within  the  tractus  olfactorius.  During  their 
course  they  give  off  collaterals,  which  end  in  free  branches  within  the  superficial  and  deep 
strata  of  the  molecular  layer. 

The  brush-cells  are  often  spindle-form  in  shape  and  horizontally  placed.  The  larger 
cells  are  found  within  the  molecular  layer,  external  to  the  mitral  cells,  which  latter  they 
in  general  resemble  in  giving  off  the  two  kinds  of  dendrites  and  in  sending  their  nerve- 
processes  to  the  granule-layer.  The  small  brush-cells,  also  known  as  the  peripheral  brush- 
cells,  lie  close  beneath  and  between  the  glomeruli.  They  likewise  send  one  dendrite  to 
the  glomerulus,  the  nerve-processes  behaving  like  those  of  the  large  brush-cells. 

5.  The  granule-layer.  Within  this  stratum  are  found  the  granule  cells  or 
granula,  peculiar  small  elements  with  long  processes.  These  granula  also  penetrate 
between  the  mitral  cells  and,  beyond  these,  into  the  molecular  layer  as  far  as  the  glom- 
eruli. The  granule  cells  are  triangular,  resembling  the  pyramidal  cells,  or  fusiform  or 
pear-shaped,  all  being  placed  vertically.  An  outwardly  directed  process,  mostly  single 
but  rarely  double,  divides  repeatedly  after  a  longer  or  shorter  course,  usually  close 
beneath  the  mitral  cells,  to  form  a  brush-like  terminal  arborization,  that  ends  within  the 
most  superficial  region  of  the  molecular  layer  at  the  glomeruli  in  delicate  filaments..  In 
the  opposite  direction,  that  is  inward,  the  granules  exhibit  several  processes,  which  are 
usually  smooth  and  slightly  branched  and  end  free  after  a  short  course.  As  yet,  a 
nerve-process  has  not  been  discovered.  In  addition  to  the  granules,  this  layer  contains 
cells  of  Golgi's  II  type — multipolar  elements  with  fusiform  or  polygonal  cell-body  and 
a  nerve-process  that  breaks  up  within  the  granule-layer.  The  nerve-fibres  running  within 
the  granule-layer  are  partly  the  nerve-processes  of  the  mitral  and  brush  cells;  further, 
fibres  enter  the  bulbus  to  end  some  within  the  granule-layer,  and  some  within  the  molec- 
ular layer  in  the  vicinity  of  the  glomeruli,  after  having  penetrated  the  layer  of  mitral  cells. 

The  nerve-processes  of  the  mitral  and  brush  cells,  that  pass  to  the  tractus  olfacto- 
rius, end  within  the  cortex  of  the  tractus  and  of  the  tuberculum  olfactorium  and  within 
the  olfactory  area  of  the  substantia  perforata  anterior  and  the  adjoining  parts  of  the  sep- 
tum pellucidum.  These  end-stations  exhibit  the  structure  of  a  modified  cerebral  cortex. 

GYRUS  FORNICATUS. 

The  structure  of  the  cortex  of  the  gyrus  fornicatus  deviates  from  the  typical  make-up 
of  the  cerebral  cortex  chiefly  in  relation  to  the  layer  of  the  large  pyramidal  cells.  Within 
the  gyrus  cinguli,  this  stratum  contains,  in  the  outer  half,  few  small  pyramid  cells  and, 
in  the  inner  half,  those  of  medium  size.  The  latter,  almost  all  of  uniform  diameter, 
lie  deeply  placed  and  together,  in  consequence  of  which  disposition  the  middle  portion 
of  the  layer  appears  poor  in  cells  and,  on  account  of  the  ascending  primordial  branches 
.  of  the  pyramidal  cells,  is  called  the  stratum  radiatum.  Towards  the  corpus  callosum, 
all  layers  suffer  diminution  in  thickness  and  in  the  size  of  the  cells.  The  cortex  of  the 
gyrus  hippocampi  bears,  in  many  respects,  a  close  resemblance  to  that  of  the  gyrus  cin- 
guli. That  part  of  the  gyrus  hippocampi  which  borders  the  fissura  collateralis  and  rhinica, 


120 


THE   FIBRE-TRACTS. 


however,  exhibits  slight  deviation  from  the  common  type.  Towards  the  fissura  hippo- 
campi, the  molecular  layer  is  broader.  Within  the  layer  of  small  pyramidal  cells,  the 
cells  are  irregularly  arranged  in  chains  of  hillocks,  while  within  the  third  layer  are  found 
larger  pyramidal  cells  with  very  long  primordial  branches  ;  moreover,  of  these  cells  the 
largest  are  very  deeply  placed,  whereby  the  conspicuous  radial  striation,  the_  stratum 
radiatum,  is  produced.  The  layer  of  polymorphic  cells  contains  almost  exclusively  small 
irregular  cells,  that  are  embedded  within  a  close  network  of  nerve-fibres. 

HIPPOCAMPUS  AND  GYRUS  DENTATUS. 

The  hippocampus,  or  cornu  Ammonis,  and  the  gyrus  dentatus  represent  two  spe- 
cial convolutions.  On  following  the  gyrus  hippocampi  dorsally,  one  reaches  the  subin- 
culum,  that  constitutes  that  part  of  the  hippocampal  convolution  in  which  gradually 
begins  a  change  in  the  structure  of  the  cerebral  cortex,  leading  finally  to  the  typical 
structure  of  the  hippocampus.  The  white  substance  splits  into  two  layers  :  the  one 
passes  to  the  free  surface  of  the  hippocampus  and  constitutes  the  alveus,  the  other  passes 
to  the  lateral  wall  and  roof  of  the  inferior  horn  of  the  lateral  ventricle.  The  alveus  is 


Layer  of 
polymorphic  cells 


Layer  of 
pyramidal  cells 

Molecular  layer 

FIG.  117. — Hippocampus  or  cornu  Ammonis  and  gyrus  dentatus  in  transverse  section.     Schematic. 


THE   HIPPOCAMPUS.  121 

continuous  with  the  fimbria.  The  uppermost  layer  of  the  gray  substance,  the  substantia 
reticularis  alba,  corresponding  to  the  molecular  layer  of  the  typical  cortex,  divides  into 
a  superficial  and  a  deep  stratum.  The  superficial  layer  adjoins  the  molecular  layer  of  the 
gyrus  dentatus  and  forms  the  lamina  medzillaris  circumvoluta.  The  deep  layer  forms  the 
stratum  lacunosum,  that  arches  around  and  embraces  the  lamina  medullaris  and  ends  in  a 
recurved  hook  at  the  medial  side  of  the  cell-layer  of  the  gyrus  dentatus.  Between  the 
lamina  medullaris  circumvoluta  and  the  stratum  lacunosum,  lies  the  stratum  moleculare. 
The  pyramidal  cells  of  the  subinculum  gradually  collect  into  a  single  layer  of  cells  as  they 
approach  the  hippocampus.  At  first  the  arrangement  of  the  cells  is  still  irregular,  towards 
the  gyrus  dentatus  the  cells  form  a  single  thick'  layer,  but  within  the  terminal  sheet  of 
the  hippocampus  they  once  more  are  irregularly  disposed.  In  this  way  two  special  zones 
are  produced,  a  deep  layer  of  pyramidal  cells,  the  stratum  lucidum,  and,  between  the 
latter  and  the  stratum  lacunosum,  the  stratum  radiatum,  so  called  on  account  of  the 
traversing  long  primordial  stems  of  the  pyramidal  cells.  The  layer  of  polymorphic  cells 
is  known  as  the  stratum  oriens.  The  gyrus  dentatus  exhibits  three  strata :  the  molec- 
ular layer,  the  granule-layer  or  the  stratum  granulosum,  and  the  layer  of  polymorphic 
cells.  The  relations  between  the  strata  of  the  modified  cortex  of  the  special  convolutions 
and  those  of  the  typical  cerebral  cortex  are  shown  in  the  following  table  and  Fig.  117: 


Cerebral  Cortex 
Molecular  layer 

Layer  of  pyramidal  cells 

Layer  of  polymorphic 
cells. 

White  substance 

Hippocampus 

f  Lamina  medullaris  circumvoluta 
<  Stratum  moleculare 
I  Stratum  lacunosum 
(  Stratum  radiatum 
|  Stratum  lucidum 

<  Stratum  oriens 
N  Alveus 

Gyrus  dentatus 

V  Molecular  layer 

1  Granule-layer   or   stratum 
granulosum 

1  Layer  of  polymorphic  cells 
or  stratum  oriens 

HIPPOCAMPUS. 
The  individual  layers  exhibit  the  following  cells: 

1.  Lamina  medullaris  and  stratum  moleculare; 

a.  Small  cells  of  Golgi  II  type. 

b.  Fusiform  cells,  with  nerve-processes  that  break  up  in  stratum  moleculare. 

2.  Stratum  lacunosum: 

Small    triangular    or    stellate  cells,    with    ascending    and    descending    dendrites    and 
nerve-processes  that  split  up  in  the  stratum  lacunosum. 

3.  Stratum  radiatum  : 

a.  Cells    of    the    same    character    as    those    of    the    stratum    lacunosum — aberrant 

cells  of  the  stratum  lacunosum. 

b.  Pyramidal  cells — aberrant  cells  of  the  stratum  lucidum. 

c.  Cells  of  Golgi  II  type. 

d.  Triangular   or   fusiform    cells,    descending    nerve-processes    ending   around    the 

pyramidal  cells. 


122  THE   FIBRE-TRACTS. 

4.  Stratum  lucidum: 

Pyramidal  cells,  with  long  primordial  stems  ascending  within  the  stratum  radiatum 
and  nerve-processes  running  to  the  alveus.  Within  the  portion  of  the  hippo- 
campus bordering  the  gyrus  dentatus,  giant  pyramidal  cells  are  found. 
Each  of  the  nerve-processes  of  these  elements  gives  off,  soon  after  its  origin 
from  the  cell,  a  collateral,  which  traverses  the  stratum  radiatum  and  passes 
to  the  stratum  lacunosum. 

5.  Stratum  oriens: 

a.  Aberrant  pyramidal  cells. 

b.  Cells  with  ascending  nerve-processes,  that  end  around  the  pyramidal  cells. 

c.  Martinotti  cells. 

Along  with  the  fibres  passing  from  the  cortex  to  the  alveus,  are  found  also  those 
which  come  from  the  alveus  and  end  within  the  cortex.  * 

GYRUS  DENTATUS. 

The  gyrus  dentatus  constitutes  a  small  modified  cerebral  cortex,  which  adjoins  the 
lamina  medullaris  circumvoluta  of  the  hippocampus  and  receives  within  its  hilus  the  end 
of  the  hippocampus.  The  white  substance  of  the  gyrus  dentatus  is  net  directly  applied 
to  the  layer  of  polymorphic  cells,  but  is  separated  from  the  latter  by  the  cortical  forma- 
tion, which  corresponds  to  the  region  of  the  hippocampus  bordering  the  gyrus  dentatus. 
It  follows,  that  the  fibres  coming  from  the  gyrus  dentatus  break  through  the  end  of  the 
hippocampus  lying  within  the  hilus,  the  alveus,  therefore,  representing  the  cortical  white 
substance  of  both  the  hippocampus  and  the  gyrus  dentatus. 

Passing  from  the  fissura  hippocampi  towards  the  ventricle,  the  following  strata  are 
encountered  : 

a.  Molecular  layer,  bordering  the  lamina 

medullaris  of  the  hippocampus, 
.     „,  ,  \  Gyrus  dentatus. 

b.  Stratum  granulosum, 

c.  Layer  of  polymorphic  cells, 

d.  Molecular  layer, 

e.  Layer  of  giant  pyramidal  cells, 

,    ,  '*     .          ...        „  \   Hippocampus. 

f.  Layer  of  polymorphic  cells, 

g.  Alveus. 

The  cells  exhibited  by  the  individual  strata  of  the  gyrus  dentatus  are  the  following: 

1.  Molecular  layer: 

a.  Cells  of  Golgi  II  type, 

b.  Aberrant  granule-cells. 

2.  Stratum  granulosum. 

This  layer  is  formed  of  the  granule-cells,  closely  placed  and  disposed  in  several 
rows.  The  cells  are  modified  pyramidal  cells,  distinguished  by  the  absence  of  the  basal 
dendrites  and  a  primordial  stem.  The  ascending  dendrites  end  within  the  molecular 
layer,  while  the  nerve-process  passes  through  the  layer  of  polymorphic  cells,  then 
through  the  molecular  layer  and  the  stratum  of  pyramidal  cells  of  the  hippocampus,  and 
exhibits  during  its  further  course  peculiar  local  thickenings,  with  small  protruding  out- 


CEREBRAL   LOCALIZATION,  123 

growths.  The  nerve-processes  unite  into  a  bundle  and  then  end,  after  forming  a 
reticular  plexus,  around  the  cell-bodies  of  the  large  pyramidal  cells  and  their  dendrites. 
They  establish  relations,  therefore,  between  the  granule-cells  and  the  giant  pyramidal 
cells  of  the  hippocampus,  from  which,  in  turn,  the  impulse  may  be  conveyed  to  other 
pyramidal  cells  by  the  collaterals  that  pass  to  the  stratum  lacunosum. 

3.  Layer  of  polymorphic  cells: 

a.  Cells  with  ascending  nerve-processes,   ending  within  the  granule-layer, 

b.  Cells  with  ascending  nerve-processes,   passing  to  the  alveus, 

c.  Cells  of  Golgi  II  type. 

As  in  the  hippocampus,  so  here,  among  the  fibres  passing  from  the  gyrus  dentatus 
are  those  coming  from  the  alveus  and  ending  within  the  gyrus  dentatus. 

In  its  further  course,  the  gyrus  dentatus  continues,  as  the  induseum  griseum,  over  the 
corpus  callosum.  The  medial  and  lateral  thickenings,  the  stria  Lancisii  and  the  taenia  tecta, 
likewise  display  the  character  of  cortex  ;  within  the  stria  Lancisii  a  molecular  layer  with 
tangential  fibres,  a  middle  layer  with  fusiform  cells  and  a  deep  layer  are  distinguishable. 

CEREBRAL  LOCALIZATION. 

The  various  subdivisions  of  the  brain  are  broadly  divided,  with  regard  to  their 
functions,  into  two  chief  groups,  the  higher  and  the  lower.  The  higher  subdivisions  are 
the  cerebral  hemispheres,  and  in  these  the  cerebral  cortex,  which  through  the  great 
development  of  the  cerebral  mantle  and  the  formation  of  the  convolutions  attains  such 
extraordinary  expansion,  plays  the  principal  role  and  represents  the  material  substratum 
of  intellectual  activity.  The  lower  subdivisions  are  interposed  between  the  cerebral 
hemispheres  and  the  spinal  cord  and  include  the  medulla  oblongata,  the  pons,  the  cere- 
bellum, the  region  of  the  corpora  quadrigemina,  and  the  cerebral  ganglia — that  region, 
therefore,  also  designated  as  brain-stem.  These  lower  parts  possess  no  direct  import  for 
intellectual  activity,  but  have  rather  the  task  of  regulating,  independently  of  conscious- 
ness or  volition,  the  many  functions  necessary  for  the  maintenance  of  the  body.  "The 
lower  brain  segments  supply  an  apparatus,  by  which  the  general  condition  of  the  body 
may  be  reflected  from  within.  For  the  moulding  of  the  intellectual  processes,  the 
mechanism  of  the  cerebrum  proper  is  authoritative." — (Flechsig. ) 

It  is  unquestionably  the  service  of  the  anatomist,  Franz  Joseph  Gall,  first  to  have 
recognized  the  significance  of  the  cerebral  cortex  for  intellectual  activity.  Since  Gall, 
anatomists  have  ceased  to  seek  a  definite  point  in  the  brain,  to  which  all  motor  and 
sensory  nerves  converge  and  which  might  be  identified  anatomically  as  the  seat  of  the 
centralized  soul.  As  well  known,  Ren6  Descartes  interpreted  the  pineal  body  as  the 
organ  of  the  soul  ;  Sommering  located  the  sensorium  commune  in  the  fluid  of  the  ven- 
tricles ;  according  to  Varolius,  the  soul  had  its  seat  in  the  soft  brain-substance ;  Thomas 
Willis  regarded  the  central  ganglia  as  perception-centres,  and  the  corpus  callosum  as  the 
seat  of  the  imagination,  while  he  placed  thought  within  the  cerebral  convolutions.  Gall, 
moreover,  established  the  principle,  that  the  individual  convolutions  are  not  all  intellec- 
tually equivalent  and  in  this  fundamental  view  already  approached  the  modern  theory  of 
localization.  In  setting  up  his  own  localization  theory  he  went  too  far,  in  that  he  sub- 
divided the  entire  cerebral  surface  into  twenty-seven  separate  areas,  which  areas  were  the 


124 


THE  FIBRE-TRACTS. 


carriers  of  definite  intellectual  faculties  and,  further,  that  along  with  the  greater  develop- 
ment of  such  a  definite  brain-area,  a  corresponding  stronger  projection  appeared  on  the 
skull.  Following  the  theory,  the  possibility  was  assumed,  that,  by  careful  examination 
of  the  skull,  a  person's  endowment  or  character  might  be  determined.  In  the  scientific 
world,  Gall's  phrenology  did  not  long  endure.  Although  the  present  theory  of  localiza- 
tion differs  entirely  in  its  essentials  from  phrenology,  we  must  nevertheless  admit  that 
localization  was  advanced  more  by  Gall  than  by  the  labors  and  views  of  the  physiologist, 
Flourens,  who  defended  the  theory  of  the  equivalence  of  the  parts  of  the  cerebrum. 
Gall  and  his  pupil,  Bouillaud  (1825),  had  already  learned  that  circumscribed  injury  of 
the  cerebrum  in  the  frontal  region  may  lead  to  disturbances  of  speech. 

According  to  Gall  and  Bouillaud,  in  1836  the  French  physician,  Marc  Dax,  furnished 
the  proof,  that  motor  aphasia  appeared  only  after  disease  of  the  left  cerebral  hemisphere, 
and  in  1861  Broca  announced  the  theorem,  that  particularly  the  left  third  frontal  convo- 
lution was  the  seat  of  speech  ;  hence  this  region  is  even  to-day  commonly  called  Broca' s 
convolution. 

This  discovery  of  the  motor  speech-centre  by  Broca  was  the  foundation  of  the  theory 
of  localization.  Although  the  proof  of  the  functional  variation  of  the  cerebral  cortex 
shattered  Flourens'  teaching  of  functional  equivalence,  this  theory  was  finally  entirely 
overthrown,  not  only  by  further  pathological  experience,  but  especially  by  experimental 
physiology,  since  in  1870  Fritsch  and  Witzig  discovered  the  electrical  irritability  of  the 
cerebral  cortex.  These  investigators  succeeded  in  inducing  movements  of  certain  parts  of 
the  body  by  stimulation  of  definite  cortical  regions  by  means  of  the  galvanic  current,  and, 
further,  reached  the  conclusion,  that  while  stimulation  of  certain  cortical  regions  produced 
movements,  no  such  result  followed  stimulation  of  other  regions.  These  investigations 
were,  confirmed  and  supplemented  in  1873  by  those  of  Ferrier,  who  employed  the 
faradic  current  instead  of  the  galvanic.  In  consequence  of  these  observations,  it  was 
possible  to  establish  a  definite  cortical  region  as  the  centre  for  movement.  Other  investi- 
gators, particularly  Nothnagel,  Carville  and  Duret,  Goltz  and  Munk,  subsequently 
succeeded,  reversing  the  order,  in  producing  paralysis  of  certain  muscles  and  impairment 
of  certain  sensory  activities  by  removal  or  destruction  of  definite  cortical  regions.  These 
labors,  along  with  the  numerous  investigations  of  other  workers,  have  established  with 
increasing  stability  the  localization  of  the  functions  of  the  cerebral  cortex. 

It  is  admitted,  therefore,  that  the  individual  regions  of  the  cerebral  surface  are  not 
equivalent,  but  of  entirely  different  significance.  Each  cortical  field  presiding  over  a 
definite  function  is  designated  as  a  centre,  and  of  such  cerebral  cortical  centres,  although 
as  yet  not  accurately  delimited,  we  recognize  the  following. 

THE   MOTOR   CENTRE. 

According  to  the  newer  investigations,  the  motor  centre  embraces  especially  the 
anterior  central  or  precentral  convolution,  and,  further,  the  posterior  part  of  the  frontal 
lobe  and  the  lobulus  paracentralis.  It  includes  the  following  regions. 

a.  Upper  region  :  lobulus  paracentralis  and  the  upper  quarter  of  the  precentral 
convolution — the  centre  for  the  movements  of  the  lower  extremity.  A  further  separation 
into  particular  centres  for  certain  groups  of  muscles  is  often  made  ;  the  data,  however, 
are  so  far  from  accord,  that  a  subdivision  into  definite  subcentres  may  be  here  omitted. 


CEREBRAL   LOCALIZATION. 


125 


The  largest  part  of  the  superior  frontal  convolution,  especially  the  region  bordering  the 
paracentral  lobule  and  the  upper  fourth  of  the  precentral  convolution,  constitutes  the 
centre  for  the  muscles  of  the  trunk. 

b.  Middle    region :    the    middle    two-fourths    of    the    precentral    convolution — the 
centre  for  the  movements  of    the  upper  extremity.       The  further   delimitation  within   this 
centre  of  subcentres   for   movements  of   the 

fingers,  the  hand,  the  arms  and  the  shoulder, 
is  so  ordered,  that  the  centre  for  the  fingers 
occupies  the  lowest  position,  and  that  for 
the  shoulder  the  highest. 

c.  Lower   region  :   the  lower  fourth 
of  the  precentral  convolution — the  centre  for 
the    musculature    of    the    face,    the    tongue, 
mastication,    the    larynx    and    the    pharynx. 
Small    special    centres    for    the    upper    and 
lower  facial  nerve  are  assumed  to  exist. 

Within  the  posterior  part  of  the  middle 
frontal  convolution  lies  the  centre  for  the 
movements  of  the  eyes  and  of  the  head, 
particularly  for  the  direction  of  the  head  and 

the  eyes  towards  the  opposite  side  (conjugate  deviation).  According  to  other  investi- 
gators, a  second  projection  centre  for  the  winking  movements  of  both  eyes  has  its  seat 
in  the  gyrus  angularis. 


FIG. 


. —  Cerebral    localization, 
centres. 


Motor     and     auditory 


FIG.  119. — Cerebral  localization.     The  chief  regions  of  the  motor  centre. 

In  regard  to  the  motor  centres  it  is  particularly  to  be  noted,  that  stimulation  within  the 
centre  calls  forth  contraction  and  movements  of  the  corresponding  muscle-area  of  the  opposite 
half  of  the  body,  and  that  in  like  manner  injuries  lead  to  paralysis  in  the  opposite  side  of  the 
body.  This  will  be  further  discussed  in  connection  with  the  motor  conducting  paths  (page  139). 


126 


THE   FIBRE-TRACTS. 


This  rule  is  not,  however,  without  exception.  From  certain  centres,  not  only  the 
corresponding  muscles  of  the  opposite  side  are  controlled,  but  also  those  of  the  same 
side;  that  is,  there  exists  for  certain  muscles  a  bilateral  innervation.  This  is  true  for 
those  muscles  whose  action,  as  a  rule,  is  not  unilateral  but  symmetrically  bilateral,  as,  for 
example,  in  the  case  of  the  frontalis,  orbicularis  oculi  and  corrugator  supercilii  muscles 
supplied  by  the  upper  facial  branch,  or  the  muscles  of  mastication,  of  the  pharynx  and 
of  the  larynx.  This  bilateral  innervation  explains  the  fact,  that  after  unilateral  destruction 
of  such  centres  the  paresis  of  the  muscles  concerned  is  not  pronounced,  since  the 
necessary  stimulus  may  still  be  supplied  from  the  unaffected  centres  of  the  opposite 
hemisphere. 

THE   SENSORY   CENTRES. 

a.  The    sensory  area,  including  the  centres  for  touch,   pain  and   temperate  sensi- 
bility, embraces  especially  the  postcentral  convolution   and  the   adjoining  anterior  part  of 
the   parietal    lobe   and,    perhaps,    even    extends    onto    the    precentral    convolution.       The 
position-  and    movement-sensibility,  as  well    as    space-  and    orientation-sense,    are   trans- 
ferred to  this  same  region.      The  impulses,  passing  to  the  sensory  area,   come  essentially 
from  the  opposite  half  of  the  body. 

b.  The   auditory   centre    is    located   in   the    middle    part  of  the-gyrus  temporalis 
superior  and  includes  additionally  the  gyri   transversi  of  the  upper  temporal  convolution, 

that    lie    buried    within    the 
Motor  centre  fissure  ccrebri  lateralis. 

c.  The  visual  centre 
lies   within   the  cuneus,   par- 
ticularly within  the  cortex  of 
the  fissura  calcarina,  perhaps 
extending     onto     the    gyrus 
lingualis. 

d.  The  olfactory  cen- 
tre   is    situated    within    the 
anterior   part    of    the    gyrus 
hippocampi   and    the    hippo- 

FIG.  120. — Cerebral  localization.    Visual  and  olfactory  centres.  CampUS. 

e.  The  gustatory  cen- 
tre has  as  yet  not  been  definitely  located,  but  probably  adjoins  that  for  smell. 

The  motor  centres  and  the  individual  sensory  regions  or  sense-centres  are 
designated  also  as  projection  centres,  for  the  reason  that  the  impulses  passing 
from  the  stimulus-receiving  organs  (skin,  muscles;  joints  and  the  higher  sense- 
organs),  and  conveyed  by  the  sensory  nerves  of  the  central  nervous  system, 
are  radiated  or  projected,  as  it  were,  to  the  sense-centres,  while  the  impulses  from 
the  motor  centres  are  similarly  projected  towards  the  periphery  and  conveyed  by  the 
motor  nerves  especially  to  the  muscles.  Stimulation  within  the  sense-centres  induces 
sensation  (touch,  sight,  hearing,  smell,  etc.)  ;  stimulation  within  the  motor  zone 
leads  to  movements.  These  incoming  and  outgoing  conductions  follow  quite  definite 
paths,  which  are  termed  afferent  or  centripetal  and  efferent  or  centrifugal  projection- 
tracts  respectively. 


ASSOCIATION   CENTRES.  127 

Viewing  the  surface  of  the  cerebral  hemispheres  and  imagining  the  individual  pro- 
jection centres  outlined,  it  will  be  evident  that  the  latter  occupy  only  a  certain  part, 
perhaps  a  third,  of  the  entire  cerebral  cortex.  In  addition  to  these  motor  and  sensory 
fields,  there  remains  a  large  area  that  embraces  certain  parts  of  the  frontal,  parietal, 
occipital  and  temporal  lobes,  together  with  the  deeply  situated  central  lobe  or  the 
insula — a  tract  of  still  slightly  known  function.  According  to  Flechsig,  this  entire  large 
area  is  designated  the  association  centres,  of  which,  following  him,  an  anterior,  a  middle 
and  a  posterior  association  centre  are  distinguished.  The  anterior  or  frontal  centre  in- 
cludes the  fore-part  of  the  frontal  lobe;  the  middle  or  insular  centre,  the  island  of  Reil; 
and  the  posterior  or  parieto-occipito-temporal  centre,  a  large  part  of  the  occipital  and 
temporal  lobes  and  almost  the  entire  parietal  lobe. 

According  to  the  theory  advanced  first  by  Flechsig,  these  association  areas 
constitute  the  substratum  for  the  higher  psychic  functions — an  apparatus  which  collects 
the  activities  of  the  sense-centres  to  higher  unity,  and  comprises  centres  of  all  the  more 
complex  associations.  They  are  the  chief  bearers  of  what  we  call  experience,  knowledge 
and  cognizance  and,  in  part,  of  speech,  in  short  the  intellectual  centres  proper.  Flechsig 
was  led  to  the  advancement  of  this  theory  chiefly  through  his  histological  investigations 
based  on  the  method  of  the  development  of  the  medullary  sheath.  He  proved  that  the 
myelin-ripening  of  the  individual  nerve-tracts  proceeds  from  below  upward,  that  is  from 
the  spinal  cord  and  the  lower  brain-segments  towards  the  cortex  of  the  end-brain. 
Already  at  birth,  according  to  Flechsig,  the  individual  tracts  have  reached,  in  large 
part,  their  development  wichin  the  lower  divisions  of  the  brain,  while  within  the  cerebrum 
only  few  paths  of  conduction  have  developed.  At  first,  one  sense-conductor  after  the 
other  gradually  pushes  out  towards  the  cerebral  cortex.  In  the  new-born  child,  only 
two  of  the  sense-centres,  the  olfactory  and  gustatory,  are  developed ;  then  follow  the 
centres  for  tactile  sense,  for  sight  and,  lastly,  for  hearing.  Only  subsequent  to  the  com- 
pleted development  of  the  sense-centres,  does  the  development  of  the  intellectual  centres 
begin  within  the  individual  territories.  Medullary  fibres  proceed  from  the  projection 
centres  to  the  neighboring  association  areas,  the  latter  likewise  become  functionally  active 
and  eventually  numerous  tracts  bind  both  kinds  of  centres  with  each  other.  Based  on 
further  investigations,  Flechsig  later  subdivided  the  entire  cerebral  cortex  into  thirty-six 
different  areas,  according  to  the  time  of  completed  myelination.  The  areas  first  be- 
coming medullated  correspond  to  the  projection  centres ;  then  follow  the  embryonic 
intermediate  centres  and,  finally,  the  terminal  districts,  which  exclusively  form  the 
association  centres. 

According  to  Flechsig,  the  projection  centres  differ  also  anatomically  from  the 
association  ones,  since  only  the  former  are  connected  by  centripetal  and  centrifugal  pro- 
jection tracts  with  the  lower  brain-centres,  while  within  the  association  centres  such  projec- 
tion tracts  are  altogether  wanting.  The  association  areas  are  connected  by  fibre-tracts 
only  with  the  projection  centres,  trom  which  they  receive  sensory  stimuli ;  on  the  other 
hand,  they  may  influence  the  sensory  areas  by  reflex  stimuli  or  inhibition.  The  associa- 
tion and  projection  centres  also  vary  in  their  histological  make-up,  since  the  association 
centres  exhibit  a  specific  although  uniform  texture,  while  the  projection  centres  present  a 
structure  which  not  only  differs  from  that  of  the  association  centres,  but  varies  within  the 
individual  fields. 


128  THE   FIBRE-TRACTS. 

This  important  theory  of  Flechsig,  however,  can  no  longer  be  accepted  in  its 
entirety.  Further  investigations  have  not  substantiated  the  assumption,  that  only  a  por- 
tion of  the  cortex  is  connected  with  the  lower  lying  brain-centres  by  means  of  projection 
tracts,  since  such  paths  have  been  proven  also  for  the  association  fields  mapped  out  by 
Flechsig.  Furthermore,  it  has  been  determined,  that  not  only  the  projection  centres  possess 
a  special  and  for  each  region  specific  texture,  but  that  there  also  exists  within  the  asso- 
ciation tract  a  large  number  of  areas  of  different  structure.  As  has  been  shown  by  the 
investigations  of  Vogt  and  Brodmann,  the  entire  cerebral  cortex  may  be  mapped  out  in 
numerous  fields,  which  differ  from  one  another  in  regard  to  cellular  stratification,  as  well 
as  in  regard  to  fibre-relation,  there  existing  a  cyto-  and  a  myelo-architectonic  differentia- 
tion of  the  cerebral  cortex  (Figs.  114  and  115). 

Concerning  the  relations  of  each  individual  -anatomically  definable  field  to  function, 
however,  we  still  know  very  little,  and  it  remains  for  future  physiological  and  clinico- 
pathological  investigations  to  advance  our  understanding  concerning  this  problem.  With 
Brodmann  we  may  assume,  ' '  that  each  specific  cytological  difference  must  be  the  expres- 
sion of  a  definite  physiological  dignity  and  that,  therefore,  all  the  variously  structured 
cortical  fields  also  preside  over  different  functions.  Not,  of  course,  in  the  sense  that  one 
assigns  complex  intellectual  processes  or  attributes  to  specially  delimited  territories,  but  in 
the  only  warranted  sense  of  Wernicke,  who  associates  only  the  most  elementary  functions 
with  definite  localities  of  the  cerebral  cortex."  A  question,  for  which  the  answer  has  long 
been  sought,  is  the  existence  of  definite  recollection  or  memory  centres.  Many  facts 
point  to  the  actual  existence  of  such  memory  centres  beside  the  projection  centres.  Thus, 
clinical  cases  are  known,  in  which  loss  of  a  perception  region  was  attended  with  cessation 
of  the  corresponding  perception,  but  not  of  the  related  memory-pictures ;  on  the  contrary, 
certain  cases  with  cortical  lesions  in  the  immediate  vicinity  of  the  perception  centres,  for 
example,  of  the  convolutions  adjoining  the  visual  and  auditory  centres,  exhibited  neither 
blindness  nor  deafness,  but  failure  of  memory  and  disturbances  of  the  function  of  recog- 
nition. Thus  lesions  of  both  occipital  lobes  lead  to  so-called  visual  agnosia  or  perception 
blindness.  The  patient  may  still  be  able  to  give  information  regarding  the  form  and 
color  of  an  object,  but  the  object  itself  is  unknown,  he  being  no  longer  able  to  recog- 
nize the  object  or,  usually,  its  spatial  relations.  Further,  lesions  of  the  left  temporal  lobe 
cause  the  so-called  auditory  agnosia  or  perception  deafness,  in  which  condition  not  only 
the  spoken  words,  but  also  auditory  stimuli  of  all  kinds  are  no  longer  understood. 
Likewise,  lesions  situated  in  the  middle  third  of  the  postcentral  convolution,  or  farther 
backward  in  the  parietal  lobe,  may  lead  to  so-called  tactile  agnosia,  in  which,  for 
example,  the  form  of  any  object  no  longer  is  recognized,  notwithstanding  the  integrity 
of  the  individual  impressions  necessary  for  touch-,  space-  and  muscle-sense. 

THE   SPEECH    CENTRES. 

The  speech  centre  in  its  entirety  includes  certain  cortical  areas  of  the  lateral  surface 
of  the  hemisphere  and,  in  right-handed  individuals,  is  located  on  the  left  side. 

a.  The  motor  speech  centre,  Broca's  centre  presiding  over  the  ability  to  speak, 
lies  within  Broca's  convolution  embracing  the  base  of  the  gyrus  frontalis  inferior.  It 
extends,  perhaps,  also  to  the  adjacent  part  of  the  lowest  region  of  the  precentral  convo- 
lution and  to  the  anterior  part  of  the  insula.  Upon  the  integrity  of  this  centre  depends 


THE   SPEECH    CENTRES. 


129 


PIG.  121. — Cerebral  localizati 


Speech  centre. 


the  ability  to  carry  out  the  co-ordinated  movements  necessary  in  speaking.  Damage  of 
the  centre  leads,  therefore,  to  abolition  of  the  execution  of  motor  speech.  Voluntary 
speech,  repeating  words  or  reading  aloud  are  no  longer  possible.  This  centre,  therefore, 
is  also  termed  the  centre  of  motor  aphasia. 

b.  The    sensory    speech    centre,  the   tone-picture  or  auditory  centre,   lies  within 
the  posterior  third  of  the  gyrus  temporalis  superior  and  the  adjoining  part  of  the  gyrus 
supramarginalis.      It  is  also  known  as   Wernicke1  s  centre  and  represents  the  cortical  region 
where  the  memory-pictures  of   the  heard  and  spoken  words  are  retained.     If    the  centre 
be  injured,  the  patient,  while 

Still        hearing       the       Spoken  .  Auditory  centre  -  Wernicke 

word,  can  no  longer  compre- 
hend what  he  hears.  He  has 
lost  speech-understanding. 
The  centre  is  also  designated 
as  the  centre  for  word-deaf- 
ness or  sensory  aphasia. 

c.  The   visual    cen- 
tre, where  the  memory-pic- 
tures  of  written    characters 
are  stored,  lies  within   the 
gyrus  angularis.     Injury  of 

the  centre  is  followed  by  inability  to  recognize  the  printed  or  written  letters,  or  to  form 
words  from  them.  The  centre  is  also  termed  the  centre  for  word-blindness  or  alexia. 

d.  A  special  writing  centre  is  still  often  assumed  to  lie  within  the  base  of  the  gyrus 
frontalis  medius.       Its   existence,   however,    is   scarcely   longer  to   be   accepted,  since  the 
centre  for  writing  is  blended  with  the  motor  centre  for  the  hand  within  the  middle  region 
of  the  precentral  convolution. 

These  speech  centres  are,  therefore,  centres  of  memory,  namely,  for  the  movement- 
conceptions  of  articulation,  the  acoustic  pictures  of  speech  and  the  visual  pictures  of 
written  speech.  Individuals,  who,  in  consequence  of  lesions  of  these  centres,  have  lost 
the  memory  of  the  motor,  auditory  and  visual  conceptions  of  speech,  are  neither  paralyzed, 
deaf,  nor  blind,  but  only  wanting  in  speech-understanding.  We  are,  therefore,  warranted 
in  assuming  the  existence  of  memory  centres  beside  the  projection  centres  ;  further,  it 
must  be  noted,  that  the  projection  centres  serve  not  only  sensation  and  innervation,  but 
also  memory,  and,  on  the  other  hand,  that  the  regions  adjoining  the  projection  centres 
are  not  to  be  regarded  as  exclusive  commemorative  centres,  since  the  existence  of 
projection  tracts  to  them  has  been  proven. 

Concerning  the  association  function  of  the  cerebrum,  we  must  assume  that  the 
binding  together  of  conceptions  or  recollections  of  the  same  kind  occurs  in  the  cortex 
within  the  individual  cortical  fields,  but  that  all  the  higher  association  processes  are  con- 
nected with  the  collective  activity  of  many,  perhaps  of  all,  the  cortical  regions. 

Finally,  it  must  be  especially  emphasized,  that  the  two  cerebral  hemispheres  are 
functionally  by  no  means  identical.  In  connection  with  localization  of  the  speech  centre, 
it  has  been  pointed  out,  that  in  right-handed  individuals  the  left  hemisphere  is  concerned. 
Not  only  for  speech  does  the  left  hemisphere  outweigh  the  right,  but  also  for  manipu- 

9 


1 3o 


THE   FIBRE-TRACTS. 


lation.  For  proof  of  this  we  are  indebted  especially  to  the  investigations  of  Liepmann, 
who  has  made  us  acquainted  with  the  clinical  picture  of  apraxia.  By  apraxia  is  under- 
stood the  inability  to  execute  the  appropriate  movements  during  continued  motion  ;  that 
is,  the  apraxic  patient  is  still  able  to  carry  out  certain  simple  movements,  as  flexing,  lower- 
ing, raising  or  extending  the  arm,  but  has  lost  the  ability  to  perform  combinations  of 
consecutive  movements,  such  as  made  in  greeting,  beckoning,  or  threatening.  Such 
expressive  movements  are  executed  in  an  entirely  abnormal  manner,  likewise  the  imitation 


Lejt  hand 


Right  hand 


FIG.   122. — A  lesion  in  a  causes  paralysis  on  the  right  side  and  dyspraxia,  or  impaired  combination- movement,  on  the  left 
lesions  in  b  and  c  cause  dyspraxia  on  the  left  side;  lesion  in  d  causes  paralysis  on  the  right  side. 

of  definite  movements,  and  objects  are  no  longer  properly  used  or  handled.  In  many 
lesions  of  the  left  hemisphere,  followed  by  paralysis  and  apraxia  of  the  right  hand,  a 
similar  affection  of  the  left  hand  may  be  recognized.  Moreover,  in  numerous  cases  of 
extensive  lesion  of  the  corpus  callosum,  dyspraxia  of  the  left  hand  is  present.  According 
to  Liepmann,  one  is,  therefore,  warranted  in  assuming  that  the  recollection  of  certain 
acquired  dexterities  and  also  the  'supervision  of  the  execution  of  the  same  are  in  predom- 
inating degree  concerns  of  the  left  hemisphere,  which  are  conveyed  to  the  right 
hemisphere  by  means  of  the  corpus  callosum. 


PATHS   OF  CONDUCTION.  131 


GENERAL    DIVISION   OF   THE   CONDUCTION    PATHS. 

The  entire  nervous  system  is  built  up  of  nervous  units  or  neurones.  With  regard 
to  their  physiological  tasks,  the  neurones  may  be  divided  into  two  chief  groups,  those 
which  conduct  impulses  centrifugally  and  those  which  conduct  centripetally. 

The  centrifugal  paths  serve  to  convey  impulses  from  the  central  nervous  system  to 
peripheral  organs,  especially  to  the  organs  of  movements  or  the  muscles.  These  may, 
in  a  general  way,  also  be  called  motor  paths.  The  centripetal  paths,  on  the  contrary, 
convey  impulses  coming  from  the  periphery  to  the  central  nervous  system.  By  means 
of  them  we  receive  information  of  what  goes  on  in  nature  outside  of  our  bodies  (higher 
sense-nerves)  ;  they  bring  us,  however,  also  information  of  the  processes,  which  are 
taking  place  within  all  the  organs  of  our  own  bodies,  information  of  which  we  are  in 
part  conscious  and  in  part  unconscious,  the  latter  impulses  being  continually  active  in 
regulating  the  most  diverse  functions  of  our  bodies.  v  The  centripetal  paths  are  also,  in 
a  general  way,  designated  as  sensory  paths. 

It  is  particularly  to  be  noted,  that,  as  a  rule,  more  than  a  single  neurone  is  con- 
cerned in  the  constitution  of  the  afferent  and  efferent  paths,  and  that  these  are  made  up 
of  two,  three  or  several  neurones  in  sequence.  In  this  way,  for  example,  the  great 
cortico-muscular  paths,  by  means  of  which  voluntary  movements  of  the  musculature  of 
the  extremities  are  called  forth,  consist  of  two  neurones.  The  first  neurone  extends 
from  the  motor  cortical  centre  through  the  brain-stem  to  the  spinal  cord,  where  it  ends 
within  the  gray  substance  of  the  anterior  horn.  The  second  neurone  extends  from  the 
anterior  horn  of  the  cord  to  the  muscle.  In  like  manner,  the  sensory  path  is  composed 
of  several  neurones,  which  conduct  impulses  from  the  periphery,  as  for  example  the 
integument  of  the  leg,  through  the  peripheral  nerves,  the  spinal  cord  and  the  brain-stem 
to  the  sensory  region.  The  first  neurone  conducts  the  impulse  from  the  periphery  to  the 
spinal  cord  or  to  the  posterior  column  nuclei,  the  second  from  the  cord  or  the  nuclei  of 
the  posterior  column  to  the  thalamus,  and  the  third  neurone  arises  in  the  thalamus  and 
ends  in  the  cerebral  cortex.  Owing  to  the  insertion  of  further  neurones,  the  entire 
make-up  may  become  still  more  complicated,  in  this  manner  longer  or  indirect  paths 
being  formed  in  addition  to  the  shorter  direct  ones.  Since  the  motor  and  sensory  paths 
conduct  impulses  from  the  centre  to  the  periphery  and,  conversely,  from  the  periphery 
to  the  centre,  that  is  similarly  "project,"  these  paths  are  also  called  projection  tracts. 

Two  additional  important  connecting  links,  the  association  conduction  and  the  reflex 
conduction,  exist  between  the  motor  and  sensory  paths.  They  are  established  by  means 
-of  intercentral  tracts.  Through  the  reflex  conduction,  a  reflex  movement,  the  reflex, 
is  liberated  without  the  accompaniment  of  psychic  processes.  This  conduction  is  effected 
by  the  so-called  reflex  collaterals,  although  individual  neurones,  intercalated  between 
the  centripetal  and  centrifugal  tracts,  may  also  participate.  Let  us  take  as  an  example 
of  a  simple  reflex,  the  patellar  or  the  corneal  reflex.  The  patellar  reflex  is  manifested 
by  a  contraction  of  the  quadriceps  muscle  and  extension  of  the  leg,  in  response  to 
stimulation  of  the  sensory  nerves  in  the  quadriceps  tendon,  as  when,  for  example,  the 
tendon  is  struck  with  the  percussion  hammer  below  the  patella,  while  the  leg  is  relaxed 
and  dependent.  This  entire  phenomenon  is  carried  out  by  the  following  paths :  the 
impulse  is  carried  from  the  tendon  of  the  muscle  to  the  spinal  cord,  by  way  of  the 


132 


MiucU 


MuscU 


Skin 


FIG.  123. — Schematic  representation  of   the  physiologically  different  conductions.     Red.  centrifugal  tracts;  blue,  centripetal 

tracts;  black,  intercentral  tracts. 


PATHS   OF  CONDUCTION.  133 

spinal  ganglion,  by  the  sensory  or  afferent  nerves.  On  entering  the  spinal  cord,  the 
sensory  fibre  divides  into  an  ascending  and  a  descending  branch,  which  branches  subse- 
quently end  within  the  gray  substance  of  the  cord,  or,  as  in  the  case  of  the  ascending 
branch,  first  within  the  nuclei  of  the  posterior  column.  .  Before  dividing  into  these 
branches,  hoWever,  the  entering  sensory  fibre  gives  off  a  delicate  collateral  branch,  a 
reflex  collateral,  which  runs  to  the  anterior  horn  and  there  ends.  Similar  reflex  collaterals, 
moreover,  are  also  given  off  from  the  ascending  primary  branch,  as  shown  in  Fig. 
123,  a.  By  means  of  these  reflex  collaterals,  the  impulse  may  be  directly  transferred  to 
motor  anterior  horn-cells  and  thence  conveyed  by  motor  fibres  to  the  muscle.  In  a 
similar  manner  the  corneal  reflex  or  tactile  lid-reflex  occurs.  This  reflex  consists  in  con- 
traction of  the  M.  orbicularis  oculi  on  touching  the  integument  of  the  eyelid,  the 
conjunctiva  or  the  cornea.  The  afferent  path  is  here  the  ophthalmic  branch  of  the  N. 
trigeminus.  From  the  sensory  trigeminal  fibres  entering  the  pons,  collateral  branches 
are  given  off,  which  pass  as  reflex  collaterals  to  the  nucleus  of  the  N.  facialis.  The 
efferent  path  lies  within  the  ocular  facial. 

In  place  of  the  reflex  collaterals,  however,  individual  neurones  may  transfer  the 
impulse  from  the  sensory  to  the  motor  tracts.  For  example,  a  sensory  fibre  on  entering 
the  spinal  cord  may  transfer  the  impulse  first  to  cells,  whose  axis-cylinders  do  not  leave 
the  cord,  as  do  those  of  the  motor  anterior  horn-cells  passing  to  the  periphery,  but 
enter  the  white  substance  and  divide  into  ascending  and  descending-  branches.  These 
division-branches,  after  a  longer  or  shorter  course,  end  within  the  gray  substance  of 
cord -segments  of  higher  or  lower  levels  (Fig.  123,  £).  First  within  these  segments 
occurs  the  transference  to  true  motor  cells.  In  this  manner,  not  only  the  motor  cells 
of  the  same  level  are  impressed,  but  the  stimulus  is  carried  to  higher  and  lower  lying 
cord-segments  and,  consequently,  transferred  to  a  larger  number  of  motor  neurones. 

The  further  possibility  exists,  that  the  impulse  may  be  conducted  from  the  spinal 
cord  by  the  ascending  tracts  to  the  higher  lying  subcortical  centres  and  that  first  here 
the  transference  to  the  motor  paths  occurs.  The  resulting  movements  are  mostly  more 
complicated  than  the  simple  reflexes,  although,  as  in  the  case  of  the  latter,  they  are 
unconsciously  executed.  As  shown  in  Fig.  123,  the  impulse  may  be  conducted  through 
certain  paths  from  the  spinal  cord  to  the  cerebellum,  thence  to  the  nucleus  ruber,  and 
from  the  latter  be  carried  downward  to  the  cord  and,  finally,  to  the  muscle. 

The  second  intercentral  connection  between  the  sensory  and  motor  parts  is  fur- 
nished by  association  conduction.  By  means  of  the  latter,  a  conscious  voluntary 
action  is  rendered  possible,  by  means  of  the  system  of  association  fibres  within  the 
cerebral  hemispheres.  An  impulse  is  carried  through  the  sensory  path  as  far  as  the 
cerebral  cortex  and  here  transferred  to  cells  within  a  certain  sense-centre ;  in  these  cells, 
it  may  be  assumed,  the  material  stimulus  is  released,  which  corresponds  to  psychic 
sensation.  One  may  simply  imagine,  that  from  this  locality  the  impulse  is  transferred 
by  an  additional  neurone  directly  to  the  cells  -in  the  motor  cortical  region  and  thence 
farther  carried  by  the  motor  tract.  The  cortical  connection  is,  however,  far  more 
complex,  since  only  after  traversing  numerous  intermediate  neurones  does  the  impulse 
finally  reach  the  motor  centre  and  from  there  pass  to  the  motor  path,  since  cooperation 
of  the  various  cortical  centres  must  be  assumed  in  explanation  of  the  complex  psychic 
processes. 


THE   FIBRE-TRACTS. 


CONDUCTION  PATHS  OF  THE  TELENCEPHALON. 

We  distinguish  two  chief  kinds  of  fibre -tracts,  association  fibres  and  projec- 
tion fibres. 

The  association  fibres  serve,  firstly,  to  bind  together  neighboring  or  remote 
regions  of  one  hemisphere  and,  as  such,  may  be  designated  as  the  association  fibres 
proper,  or  association  fibres  in  the  strict  sense.  Secondly,  they  serve  to  connect  the 
regions  of  both  hemispheres  and,  in  this  capacity,  are  termed  com missural  fibres. 

The  projection  fibres  unite  the  cortex  of  the  hemispheres  with  the  lower  lying 
parts  of  the  brain  and  with  the  spinal  cord — centrifugal  or  corticofugal  tracts ;  the  pro- 
jection fibres  also  include  those  passing  in  the  opposite  direction  from  the  lower  parts 
and  ending  within  the  cortex — centripetal  or  corticopetal  tracts. 

I.    ASSOCIATION   FIBRES. 

These  are  distinguished  as  short  and  long  fibres.  The  short  fibres  unite  adjoining 
convolutions  and  are  also  called  intralobular  or  U-fibres,  or  fibrae  propriae  or  arcuatae. 
The  long  fibres  connect  the  regions  of  one  hemisphere  which  are  more  or  less  separated 
and  are  also  termed  interlobar  bundles.  The  most  important  of  these  are  : 


FIG.    124. — Association    fibres,     commissural    fibres     and 
projection   fibres. 


FIG.  125. — Association  fibres.      Fasciculus   uncinatus  and 
fasciculus  longitudinalis  superior. 


FIG.    126. — Association  fibres.      Cingulum   and   fasciculi: 
longitudinalis  inferior. 


FIG.  127. — Association   fibres.     O.  P.,  fasciculus  occipito- 
frontalis;  U-,  fasciculus  uncinatus. 


PATHS   OF  CONDUCTION.  135 

a.  The  fasciculus  uncinatus,  connecting   the  orbital   surface  of  the  frontal  lobe  with 
the  temporal  pole  and  the  anterior  part  of  the  gyri  temporales. 

b.  The  fasciculus   longitudinalis   superior   or   arcuatus,    connecting    the    operculum 
frontale  and  parietale  with  the  lobulus  parietalis  inferior,   the  occipital  lobe  and  the  pos- 
terior part  of  the  upper  and  middle  temporal  convolutions. 

c.  The  fasciculus  longitudinalis  inferior,  connecting  the  occipital  pole,  the  cuneus, 
the  gyrus  lingualis  and  fusiformis  with  the  temporal  pole. 

d.  The   cingulum,   also   called   the  fornix  periphericus ,    running   within   the   gyrus 
fornicatus,  as  association  bundles  of  the  rhinencephalon. 

e.  The  fasciculus  fronto-occipitalis  (Forel-Onufrowicz),  running  immediately  beneath 
the  corpus  callosum,  over  the  nucleus  caudatus  and  within  the  corona  radiata,  and  con- 
necting  the     frontal   with    the   occipital    lobe.     According   to    recent    investigations,    this 
bundle  is  to  be  regarded  rather  as  a  projection  fibre-system. 

f.  The  association  bundles,  which   pass   through  the  capsula   externa  and  extrema. 

II.    COMMISSURAL   FIBRES. 

These  unite  both  hemispheres  and  include  : — 

a.  The  corpus  callosum,  connecting  the  cortical  districts  of  the  pallium. 

b.  The    commissura    anterior,    connecting    the    districts    belonging    to    the    rhinen- 
cephalon. 

c.  The  commissura  hippocampi,  connecting  the  same  districts  as  the  preceding. 
The  fibres    constituting   the  corpus   callosum    connect   the   cortical    districts    of   one 

hemisphere  with  those  of  the  other  hemisphere  and  form  the  radiatio  corporis  callosi, 
which  is  subdivided  into  a  pars  frontalis,  pars  parietalis,  pars  temporalis  and  pars  occip- 
italis  (page  40).  The  commissura  anterior  includes  a  pars  anterior  or  olfactoria  and  a 
pars  posterior  or  interhemisphaerica.  The  pars  olfactoria  connects  the  lobus  olfactorius 
of  one  side  with  that  of  the  opposite  side.  The  pars  interhemisphaerica  connects  the  two 
gyri  hippocampi  with  each  other, «,  The  commissura  hippocampi,  also  termed  the  fornix 
transversus,  or  lyra  Davidis,  connects  the  two  hippocampi  with  each  other. 

III.   PROJECTION   FIBRES. 

These  unite  the  cortex  of  the  telencephalon  with  the  lower  lying  parts  of  the  brain, 
as  the  corpus  striatum,  thalamus,  regio  subthalamica,  corpora  quadrigemina,  pons  and 
medulla  oblongata,  and  with  the  spinal  cord.  They  arise  from  the  crest  of  the  convo- 
lutions and  form  collectively  the  corona  radiata.  To  them  also  belong,  as  already  noted, 
fibres  which  ascend  to  the  cortex  from  lower  parts  of  the  brain.  Short  and  long  tracts 
are  distinguished. 

A.  SHORT  TRACTS. 

i.  Fibres  passing  from  all  parts  of  the  cortex  to  the  thalamus  and,  vice  versa, 
from  the  thalamus  to  the  cortex — tractus  cortico-thalamici  and  thalamo-corticales  or  the 
peduncles  of  the  thalamus.  Such  connections  include : 

a.  The  cortex  of  the  frontal  lobe  with  the  anterior  end  of  the  thalamus ; 


136 


THE   FIBRE-TRACTS. 


b.  The   cortex   of   the   central  convolutions  and  of  the  anterior  part  of  the  parietal 
lobe  with  the  outer  and  inner  thalamic  nuclei ; 

c.  The   cortex   of   the   posterior   part  of  the  parietal  and  of  the  occipital  lobes  with 
the  pulvinar  ; 

d.  The   occipito-temporal   lobe   with   the  ventral  and  medial  parts  of  the  thalamus. 
An   important  ascending    tract    from   the   thalamus  to    the   cortex    is   the   tegmental 

tract,  or  tegmental  radiation.  The  fibres  pass  from  the  ventral  region  of  the  thala- 
mus, partly  through  the  in- 
ternal capsule  direct  to  the 
cortex  and  partly  first  through 
the  lenticular  nucleus,  sub- 
sequently joining  the  fibres 
following  the  internal  capsule. 
The  course  of  those  trav- 
ersing the  lenticular  nucleus 
is  shown  in  Fig.  1 29 ;  com- 
pare also  Fig.  154.  Fibres 
pass  also  in  the  opposite 
direction,  from  the  cortex 
to  the  ventral  part  of  the 

thalamus.  The  tegmental  path  is  also  designated  as  the  tractus  cortico-tegmentalis. 
2.  Fibres  passing  from  the  cortex  of  the  visual  centre  to  the  superior  colliculus 
and  to  the  corpus  geniculatum  laterale  and,  in  the  opposite  direction,  from  the  lateral  gen- 
culate  body  to  the  cortex.  The  corpus  geniculatum  laterale  and  the  pulvinar,  together 
with  the  superior  colliculus,  constitute  the  primary  visual  centre,  the  connection  of  these 
parts  with  the  cortical  visual  centre  within  the  occipital  lobe  forming  the  optic  radiation 
of  Gratiolet.  In  this  connection  it  is  to  be  noted,  that  the  fibres  to  the  cortex  pass  only 


Corpora 
quadrigemina 

FIG.  128. — Projection  tracts.     Stalks  of  the  thalamus.      Fibres  to  the  anterior 
and  posterior  corpora  quadrigemina  and  to  the  nucleus  ruber. 


Nucleus  ruber 


Ansa  peduncularis 


\ 


Ansa  lenticularis 


\ttd-  geniculata. 


FIG.  129. — Short  projection  tracts.  Fibres  to  thalamus,  to  nucleus  ruber  and  to  the  corpora  quadrigemina.  Fibres 
of  the  tegmental  tract,  which  proceed  from  the  thalamus  and  traverse  the  lenticular  nucleus.  On  the  right,  fibres  to  the 
nucleus  caudatus  and  to  the  putamen  of  the  lenticular  nucleus. 


PATHS   OF  CONDUCTION.  137 

from  the  corpus  geniculatum  laterale,  the  chief  end-station  of  the  tractus  opticus,  and 
from  the  pulvinar  thalami,  fibres  from  the  superior  colliculus  to  the  cortex  not  being 
authenticated. 

3.  Fibres  passing  from   the  cortex  of  the  auditory  centre  to  the  inferior  colliculus 
and  to  the  corpus  geniculatum  mediate  and,   reversed,   from  the  latter  to  cortex.      As  in 
the  case  of  the  superior  colliculus,  so  also  in  that  of  the  inferior,  the  presence  of  a  quad- 
rigemino-cortical  tract  is  unproven. 

4.  Fibres    passing    from    the    cortex    (frontal     lobe,     regio     opercularis)     to    the 
nucleus    ruber. 

5.  The  fornix  passing,  as  the   equivalent  of   a  bundle   of   the  corona   radiata,  from 
the  hippocampus  to  the  diencephalon,   the  fibres  ending  within  the  corpus  mamillare. 

B.     LONG  TRACTS. 

The  fibres  pass  from  the  cortex  through  the  internal  capsule  to  the  crusta  or  basis 
pedunculi  cerebri,  to  end  within  the  pons,  the  medulla  oblongata  and  the  spinal  cord. 
The  chief  tracts  are  : — 

i.  The  frontal  pontile  tract.  The  fibres  arise  within  the  cortex  of  the  fron- 
tal lobe,  traverse  the  posterior  part  of  the  anterior  limb  of  the  internal  capsule 


Fronio-pontile  tract 


Cerebellum,  where 
pontili  fibres  end 


Pyramidal  deciissation 

FIG.  130. — The  long  projection  tracts. 

form  the   inner    fifth    of    the    basis    pedunculi    and    end    within    the    pons    in    the    pon- 
tile   nucleus. 

2.  The  occipito-temporal  pontile  tract.  The  fibres  arise  within  the  cortex  of  the 
occipital  and  temporal  lobes,  traverse  the  posterior  segment  of  the  internal  capsule,  form 
the  outer  fifth  of  the  basis  pedunculi  and  end  within  the  pons  in  the  pontile  nucleus. 
The  tractus  corticis  ad  pontem  further  is  joined  by  the  tractus  ponto-cerebellares,  con- 
necting the  pons  with  the  cerebellum  (Figs.  130  and  133). 


'38 


THE   FIBRE-TRACTS. 


3.  The  motor  tract.  The  fibres  arise  within  the  cortex  of  the  precentral 
convolution  and  the  paracentral  lobule,  pass  through  the  knee  and  anterior 
two-thirds  of  the  posterior  limb  of  the  internal  capsule,  form  the  middle  three- 
fifths  of  the  basis  pedunculi,  and  continue  to  the  medulla  oblongata  and  the 
spinal  cord.  The  entire  motor  tract  comprises  the  cortico-bulbar  and  cortico- spinal 
tracts  (Fig.  132). 

a.  The  cortico-bulbar  tract  or  tract  of  the  motor  cerebral  nerves.  The  origin  of 
the  fibres  is  known  for  only  the  facial  and  hypoglossal  nerves,  the  fibres  of  which 
arise  within  the  cortex  of  the  lower  part  of  the  precentral  convolution.  The  tract 


Frontal  and 

occipito-temporal 

pantile  tract 


Anterior  pyramidal  tract 


Cortico-bulbar  and 
cor tico-s final  tract 


Cerebellum,  ending  oj 
pantile  fibres 


Nuclei  of  motor  cerebral 
nerves  in  Medulla  oblongata 


Pyramidal  decussation 
Lateral  pyramidal  tract 


Sfinal  cord 
FIG.  131. — Long  projection  tracts. 


through   the   knee    of    the    internal    capsule    to    the    basis    pedunculi    cerebri   and 
ends    in    the    nuclei    of    the    motor    nerves    of    the    opposite   side. 

b.  The  cortico-spinal  tract  or  tract  of  the  motor  spinal  nerves.  The  fibres  of  this 
path,  also  known  as  the  tractus  cerebro-spinalis  or  the  pyramidal  tract,  take  origin  in 
the  cortex  of  the  lobulus  paracentralis  and  of  the  upper  and  middle  parts  of  the  motor 
region  of  the  precentral  convolution,  traverse  the  anterior  two-thirds  of  the  posterior 
limb  of  the  internal  capsule,  and  continue  through  the  basis  pedunculi  and  the  pons  to 
the  medulla  oblongata.  At  the  transition  of  the  medulla  to  the  spinal  cord,  the  fibres  of 
the  pyramidal  tract  cross  to  the  opposite  side,  forming  the  Pyramidal  decussation.  The 
latter,  however,  is  not  complete,  since  a  small  portion  of  the  fibres  continues  uncrossed 


PATHS   OF  CONDUCTION. 


139 


in  the  anterior  column  of  the  spinal  cord  as  the  fasciculus  cerebro-spinalis  anterior  or 
anterior  pyramidal  tract.  The  termination  of  these  fibres  is  within  the  anterior  horn  of 
the  spinal  cord  and,  moreover,  in  the  anterior  cornu  of  the  opposite  side,  the  fibres 
crossing  through  the  anterior  commissure.  The  larger  part  of  the  fibres  crosses  to  the  oppo- 
site side,  and  descends  in  the  lateral  column  of  the  spinal  cord  as  the  fasciculus  cerebro- 
spinalis  lateralis  or  lateral  pyramidal  tract,  to  end  in  the  anterior  horn  of  the  same  side. 


Cortlco-bnlbar  tract 


N.  glossopharyngens  and  Vagus 
(motor  part) 


Corttco-spinal  trad 


to* 


Lateral  pyramidal  tract. 


FIG.  132. — Cortico-bulbar  and  cortico-spinal  tracts. 

The  course  of  the  motor  tract  explains  the  fact,  that  movements  induced  by 
stimulation  of  the  motor  cortical  region  occur  chiefly  in  the  muscles,  of  the  opposite 
half  of  the  body,  or  that  injury  of  the  central  neurones  of  the  motor  tract  is 
followed  by  paralysis  of  the  muscles  of  the  opposite  half  of  the  body.  Such 
paralyses  of  one  side  (hemiplegia)  are  usually  caused  by  lesions  within  the  capsula 
interna,  less  frequently  by  lesions  within  the  cerebral  peduncle  or  the  pons.  Since 
the  speech-tract  takes  its  origin  within  the  left  hemisphere,  lesions  of  the  motor  paths 
within  the  left  hemisphere,  or  right-sided  hemiplegias,  are  usually  associated  with 
disturbances  of  speech. 


I40 


THE    FIBRE-TRACTS. 


Tract,  cerebro-sfinalis  ant. 


FIG.  133. — Motor  tract  (red)  and   the  frontal   and    occipito-temporal   pontile  tracts  (blue);   these  paths   are  continued  by 
the  ponto-cerebellar.  tract  (also  blue). 


PATHS   OF  CONDUCTION. 


141 


Tract,  cerebro-spinalis  ant. 


Medulla  oblongafa 
OliVa 


Medulla  otlongala 
Nuclei  funicul.posf. 


.  Tract    cerebro-spinalis  lat. 


Medulla  spinal/s 
FIG.  134.— Motor  tract  (red)  and  the  frontal  (F)  and  the  occipito-temporal  (07*)  pontile  tract  (blue). 


142 


THE   FIBRE-TRACTS. 


In  Fig.  136,  the  course  of  the  motor  tract  is  schematically  represented  to  explain 
the  most  important  forms  of  paralysis.  In  total  hemiplegia,  hemiplegia  completa  (Fig. 
136,  «),  the  destruction  of  an  entire  descending  motor  tract  from  one  hemisphere  is 
concerned.  In  such  cases,  the  lesion  usually  lies  within  one  motor  tract  somewhere 
along  the  brain-stem  between  the  internal  capsule  and  the  pyramidal  decussation  in 
the  medulla  oblongata,  since  within  this  stretch  all  the  descending  motor  fibres  are  com- 


Fronto-thalamic  fibres 


Fronto-pontile  tract 


Cortico-bnlbar  tract 
(  VII,  XII) 


.      Cortico-spinal  tract 
(arm) 


Cortico-spinal  tract 
(leg) 


.  Tegmental  tract 
(sensory  tract) 


FIG.  135. — Course  of  the  tracts  through  the  internal  capsule. 


pressed  into  a  field  of  small  area.  Most  frequently  the  lesion  is  situated  within 
the  internal  capsule  (knee  and  anterior  two-thirds  of  the  posterior  limb),  less  fre- 
quently within  the  cerebral  peduncle  and  the  pons.  If  in  a  lesion  of  the  motor 
tract  within  the  internal  capsule  the  knee  of  the  capsule  remains  uninvolved,  the 
facial  and  hypoglossal  nerves  do  not  share  in  the  paralysis,  such  condition  con- 
stituting hemiplegia  incompleta  (Fig.  136,  £).  In  case  the  lesion  be  located 
within  the  region  of  the  cerebral  peduncle,  the  emerging  fibres  of  the  oculomotor 
nerve  are  often  also  implicated.  Under  such  conditions  a  homolateral  oculomotor 
paralysis  exists  in  conjunction  with  the  crossed  hemiplegia,  the  condition  being 


PATHS  OF  CONDUCTION. 


143 


designated   as    hemiplegia    alternans    oculomotoria,   or    Weber's    paralysis  (Fig.     136,  c~). 

Hemiplegia    alternans     is    also     encountered     in    affections    of    the    pons  and    in    lesions 
within     the     range     of    the     medulla     oblongata.       Thus,     in     pontile     lesion,     paralysis 

of     the    extremities     on    one    side    occurs    with    paralysis    of    the    facial  nerve    on    the 

other — hemiplegia    alternans    facialis    or     Gubler's    paralysis    (Fig.     136,  d^).       Further 

combinations    are :    crossed     limb-palsy    with    homolateral     paralysis     of  the    abducens, 
or  crossed  hemiplegia  with  homolateral  hypoglossal  or  lingual  paralysis. 


Leg  centre 


Hypoglossus 


Facialis 


Hypoglossui 


Leg  tntiscles 


FIG.  136. — Schematic  representation  of   the  course  of  the  motor  tract,  explaining  the  most  important 
forms  of  paralysis. 

Hemiplegias  following  complete  destruction  of  the  entire  motor  cortical  region  of 
one  hemisphere  are  rare,  by  reason  of  the  large  extent  of  the  motor  centre.  Cortical 
diseases  are  more  frequently  limited  to  circumscribed  areas  and  paralyses  resulting 
from .  cortical  lesion  are  confined,  as  a  rule,  to  particular  portions  of  one-half  of 
the  body.  In  such  cases  one  speaks  of  monoplegia,  or,  more  definitely,  as 
monoplegia  cruralis,  monoplegia  brachialis,  or  monoplegia  facialis,  according  to  the 
involvement  of  the  motor  centre  for  the  leg,  arm,  or  face  respectively.  Such 
palsies  are  frequently  associated  with  sudden  seizures  of  convulsions  (cortical  or 
Jacksonian  epilepsy). 


i44  THE   FIBRE-TRACTS. 

A  lesion  of  both  pyramidal  tracts  descending  in  the  anterior  and  lateral  columns  of 
the  spinal  cord  leads  to  paraplegia  or  paralysis  of  both  upper  or  lower  extremities  (Fig. 
136,  f,  g} — paraplegia  brachialis  or  superior  and  paraplegia  cruralis  or  inferior.  In  very 
rare  cases,  the  lesion  may  involve  the  pyramidal  decussation  in  such  manner,  that  the 
fibres  for  one  extremity  are  interrupted  above  and  those  for  the  other  below  their  place 
of  crossing.  In  such  cases  hemiplegia  cruciata  results,  that  is,  paralysis  of  the  arm  on 
one  side  and  of  the  leg  on  the  other  (Fig.  136,  <?).  • 

RADIATIO  CORPORIS    STRIATI. 

The  corpus  striatum  is  divided  by  the  internal  capsule  into  two  parts,  the  nucleus 
caudatus  and  the  nucleus  lenticularis.  The  last-named  nucleus  is  subdivided  into  a  lateral 
portion,  the  putamen,  and  a  medial,  the  globus  pallidus,  the  latter,  in  turn,  exhibiting 
smaller  segments.  The  separation  of  the  lenticular  nucleus  into  the  individual  components 
is  effected  by  white  fibre-sheets,  the  laminae  medullares. 

CONNECTIONS   OF   THE   CORPUS    STRIATUM. 

a.  Fibres   arising    within    the   cortex    pass,  as    fibres  of   the  corona   radiata,  to   ttie 
nucleus  caudatus  and  the  nucleus  lenticularis. 

b.  Fibres  from  the   nucleus    caudatus   and    the  putamen  of   the   nucleus   lenticularis 
pass  to  the  thalamus  and  the  regio  subthalamica'. 

The  fibres  from  the  caudate  nucleus  traverse  the  internal  capsule  and  reach  the 
globus  pallidus,  while  those  from  the  putamen  pass  directly  to  the  globus  pallidus  and 
then  run,  together  with  those  from  the  caudate  nucleus,  to  the  thalamus — radiatio  strio- 
thalamica. 

Fibres,  which  course  ventrally  and  are  augmented  by  those  coming  from  the  globus 
pallidus,  pass  medially  along  the  base  of  the  lenticular  nucleus  to  the  subthalamic  region 
— radiatio  strio- subthalamica.  These  fibres  form  the  ansa  lenticularis  and  come  into  rela- 
tion partly  with  the  ventral  portion  of  the  thalamus^  and  partly  with  the  corpus  subtha- 
lamicum  or  corpus  Luysi  and  with  the  nucleus  ruber.  Some  fibres  pass  still  lower,  as 
far  as  the  mid-brain,  to  the  inferior  colliculi  of  the  corpora  quadrigemina  and  the 
substantia  nigra. 

The  ansa  lenticularis,  together  with  the  inferior  stalk  of  the  thalamus  which  conveys 
chiefly  fibres  from  the  temporal  lobe  to  the  ventral  and  medial  parts  of  the  thalamus, 
forms  the  ansa  peduncularis  (Fig.  129). 

FIBRE-PATHS  OF  THE  RHTNENCEPHALON. 

i.    PERIPHERAL   TRACT. 

This  extends  from  the  olfactory  mucous  membrane  to  the  bulbus  olfactorlus.  The 
impulse  is  conducted  by  the  peripheral  processes  of  the  intraepithelial  bipolar  olfactory 
cells  to  the  latter,  and  thence,  by  their  central  processes,  Jila  olfactoria,  to  the  glomeruli 
olfactorii. 


PATHS   OF  THE   RHINENCEPHALON. 


2.    CENTRAL   TRACT. 

A.  Connection  of  the  bulbus  olfactorius  with  the  primary  centres.  Within 
the  glomeruli,  the  impulse  is  transferred  to  the  olfactory  brush  of  the  mitral  and  brush 
cells ;  it  then  reaches  the  mitral  or  brush  cells  and  thence,  by  means  of  their  axones,  is 
conducted  centrally  to  the  primary  centres  (Fig.  137).  The  bulbus  olfactorius  is,  as  it 
were,  an  intercalated  ganglion,  being  the  end-station  of  the  peripheral  tract  and  the 


Ganglion  dor  sale  et  pro 
fund-urn  tegmenti 


Ganglion  inierpedit 
Commissnra  ant. 


Nucleus 

amygd.     Gyr.    Fhn- 
kippoc.    brio. 


FIG.  137. — Fibre-tracts  of  the  rhmencephalon.      Peripheral  tract:    olfactory  mucous  membrane  to  olfactory  bulb.    Central 
tract:   connection  of  the  olfactory  bulb  with  the  primary  centres. 

starting  point  of  the  central  tract.  The  primary  centres  include  the  gray  substance  of 
the  tractus  olfactorius  and  of  the  trigonum  olfactorium,  the  substantia  perforata  anterior 
and  the  adjoining  part  of  the  septum  lucidum. 

B.  Connection  of  the  primary  centres  with  the  secondary  or  cortical 
centres.  The  secondary  or  cortical  centres  are  :  the  gyrus  hippocampi,  the  hippocampus 
and  the  gyrus  dentatus.  The  connection  is  established  by  : 

a.  The  stria  olfactoria  lateralis.       The    fibres    pass  from    the    trigonum    olfactorium 
through  the  gyrus   olfactorius   lateralis  to  the  anterior  end  of   the  gyrus   hippocampi  and 
terminate  within  the  cortex  of  the  same. 

b.  The  olfactory  bundle  of  the  hippocampus  (Zuckerkandl),     The  fibres  arise  within 
the  trigonum    olfactorium    and    the    substantia  perforata    anterior,  extend    to   the   septum 
lucidum,  are  augmented  by  fibres  from  the  septum  and  then  pass  backward  through  the 
fornix  as  far  as  the  hippocampus. 

c.  The  stria  Lancisii.      The  fibres  pass  from  the  trigonum  olfactorium  as  the  stria 
olfactoria   medialis    towards   the  gyrus   subcallosus,  thence  over   the  corpus    callosum  and 
through  the  gyrus  dentatus  to  the  hippocampus  formation. 

According  to  Dejerine,  the  nucleus  amygdalae  is  also  a  cortical  centre.  With  this 
nucleus  a  fibre  bundle,  the  taenia  semicircularis ;  stands  in  close  relation.  The  fibres 


x46 


THE   FIBRE-TRACTS. 


arise  within  the  substantia  perforata  anterior  and  the  septum  lucidum,  are  augmented  by 
fibres  coming  from  the  anterior  commissure,  then  extend  convergingly  toward  the  sulcus 
mtermedius,  where  they  run  backward  between  the  nucleus  caudatus  and  the  thalamus 
and  end  within  the  nucleus  amygdalae.  During  the  ascending  anterior  course  of  the 
bundle,  fibres  are  given  off  at  right  angles  and  enter  the  thalamus  (Fig.  138). 

The  fornix  has  already  been  referred  to  as  the  corona  radiata  bundle  of  the 
hippocampal  formation.  The  fornix  fibres  arise  from  the  pyramidal  cells  of  the  hippo- 
campus and  the  polymorphic  cells  of  the  gyrus  dentatus.  They  extend,  first  as 
the  fimbria  and  then  as  the  posterior  limb  of  the  fornix,  toward  the  splenium  corporis 
callosi.  In  this  locality  fibres  pass  across  to  the  opposite  fornix  limb,  thus  forming 
the  fornix  transversus  or  the  commissura  hippocampi.  During  its  course  beneath 
the  corpus  callosum,  the  fornix  receives  accessions  from  the  striae  Lancisii  in  the 


Stria  Lancisii 


Taenia  semicirculari. 


Fasciculus  olfactorins 
hippocampi 


Tubercnfam  ant.  thala 


Septum  lucid. 
Tractus 


Fibrae  perforantes 

\  Ganglion  dorsale 

•L^  et  profund.  tegmenti 


Fornix 
Fascia  dentata 


Gangl.  interpeduncii! 
Corpus  mamillare 


Tuber  cinereu 

Trigon 

Stria  olfact.  lateral. 
Siibst.  perf.  ant. 
Nucleus  amygdalae 
Gyrus  hippocamp 


Fimbria    Fasc.  denial. 

FIG.  138. — Fibre-tracts  of  the  rhinencephalon.     Connections  of    the    primary    centres    with    the    secondary 

or  cortical  centres. 


form  of  fibres  which  pierce  the  corpus  callosum.  These  are  known  as  the  fibrae 
perforantes  and  constitute  the  fornix  longus  of  Forel.  In  addition  to  those  from 
the  striae  Lancisii,  other  fibrae  perforantes  from  the  gyrus  fornicatus  penetrate  the  cal- 
losum. The  fornix  fibres  continue  downward,  as  the  columnae-  fornicis,  behind  the 
anterior  commissure.  The  majority  of  the  fornix  fibres  terminate  within  the  corpus 
mamillare — tractus  cortico-mamillaris ;  another  part  of  the  fibres,  however,  passes  to. 
the  stria  medullaris  thalami  and  with  these  to  the  ganglion  habenulae,  as  the  tractus 
cortico-habenularis. 

Some  fibres  of  the  fornix  reach  their  end-station  by  another  route.  Such  aberrant 
fibres  branch  off  either  above  the  foramen  Monroi,  passing  in  front  of  the  anterior 
commissure,  or  at  the  level  of  the  tuber  cinereum,  and  course  to  the  corpus  mamillare 
as  the  stria  alba  tuberis  of  Lenhosse'k  (page  57). 

.   The  paths    proceeding  from    the  corpus  mamillare,  as  well    as    those    related  to  the 
ganglion  habenulae,  may  here  be  considered. 


PATHS   OF  THE   RHINENCEPHALON. 


The  corpus  mamillare  consists  of  two  nuclei  or  ganglia,  a  medial  and  a  lateral 
one.  The  medial  ganglion  forms  the  chief  portion,  while  the  lateral  ganglion  is  small 
and  arches  around  the  medial.  From  the  medial  ganglion  arises  the  fasciculus  mamillaris 
princeps,  which  extends  obliquely  upward  and  outward.  The  fibres  of  this  bundle  divide 
into  two  branches,  one  of  which  becomes  the  fasciculus  thalamo -mamillaris  or  tractus 
mamillo-lhalamicus,  the  other  the  fasciculus  tegmento-mamillaris  or  the  tractus  mamillo- 
tegmentalis  (Fig.  139). 

The  fasciculus  mamillo-thalamicus ,  or  the  bundle  of  Vicq  d'Azyr,  ends  with  freely 
separated  fibres  within  the  nucleus  anterior  thalami. 

The  fasciculus  mamillo-tegmentalis,  the  tegmental  bundle  of  the  corpus  mamillare 
of  Gudden,  passes  backward  and  enters  the  tegmentum  of  the  cerebral  peduncle.  The 

.Fasc   tegmento-mamillaris 
(Gndden's  tegmental  strand) 


Fibrae  perforantes  (Fornix 
•tongus) 


Tuberciilnm  anterius 
thalami 


Fasc.  thai, 
(  Vicq  d'Azyr's    bundle) 


Fornix  transversus 
Gangl.  dorsale  et  prof, 
tegm. 
Fasc.  long,  dorsa'. 

(Schutz) 
Fimbria 
Fascia  dentata 


Pedwnciihts  cnrf.  mamilla 
Ganglion  inter J>  eduuculare 


Fimbria 


FIG.  139. — Fibre-tracts  of   the  rhinencephalon.      Further  connections  of   the  cortical  centres.      The  fornix  and  the  system 

of  the  corpus  mamillare. 

major  part  of  the  fibres  ends  in  a  small  ganglion,  the  ganglion  profundum  tegmenti,  and 
in  the  neighboring  gray  substance  of  the  Sylvian  aqueduct,  some  fibres  branching  off  to 
the  posterior  longitudinal  fasciculus,  while  others  are  supposed  to  extend  as  far  as  the 
formatio  reticularis  of  the  pons. 

The  pedunculus  corporis  mamillaris  has  its  origin  within  the  lateral  ganglion  of  the 
mammillary  body.  The  bundle  courses  within  the  tegmentum  and  ends  in  the  ganglion 
dorsale  tegmenti  and  in  the  surrounding  gray  substance.  Fibres  are  also  described  as 
reaching  the  vicinity  of  the  medial  fillet.  The  dorsal  longitudinal  bundle  of  Schutz 
arises  within  the  dorsal  tegmental  nucleus  and  the  central  gray  substance  (Fig.  139). 

Concerning  the  course  and  destination  of  these  bundles  which  pass  from  the  corpus 
mamillare  to  the  tegmental  region,  we  are  by  no  means  sufficiently  informed.  According 
to  other  findings,  also  ascending  bundles  run  within  the  pedunculus  corporis  mamillaris; 
these  are  said  to  arise  within  the  tegmentum  from  the  ganglion  profundum,  as  well  as 
from  the  fillet-layer,  and  to  end  within  the  corpus  mamillare. 


I48 


THE    FIBRE-TRACTS. 


The  dorsal  longitudinal  bundle  of  Schiitz  (Kolliker's  dorsal  gray  longitudinal  bun- 
dle, Bechterew's  dorsal  longitudinal  bundle  of  the  central  gray  substance)  is  not  to  be 
confused  with  the  strand  commonly  designated  as  the  posterior  longitudinal  bundle.  The 
longitudinal  bundle  of  Schiitz  extends  through  the  gray  substance  of  the  entire  brain- 
stem  and  is  connected  with  the  nuclei  of  all  the  cerebral  nerves  and  many  other  ganglia. 
It  is  termed  the  fasciculus  longitudinalis  dorsalis,  while  the  ' '  posterior  longitudinal 
bundle ' '  is  designated  as  the  fasciculus  longitudinalis  medialis. 

The  majority  of  the  fibres  of  the  stria  medullaris  thalami  end  within  the  ganglion  habenulae. 

The  stria  medullaris  thalami  conveys  : 

a.  Fibres  coming  from  the  fornix — tractus  cortico-habenularis. 

b.  Fibres    coming   from    the   septum   lucidum   and   from   the   area   olfactoria — tractus 
olfacto-habenularis. 

c.  Fibres  coming   from    the    interior   of    the    thalamus — tractus   thalamo-habenularis. 


Stria  tltalami 


Tractns  cortico-habenularis    


Sept.  lucid. 


Sitbstantia  perf.  ant.  (Area  olfactoria) 


Fibrae  per  for  antes 
(Fornix  longns) 


Ganglion  habenulae 

Ganglion  dorsale  et  prof 

teg,,,enti 
FaSC.  long,  dorsal 

—  Schiitz  — 
Fimbria 


Fasc.   retroflexiis 
—  Meynert  — 
Fasc.  haben.  Gangl.  tnterped. 
Gangl.  interped. 


FIG.  140. — Fibre-tracts  of   the  rhinencephalon.      Further  connections  of   the  cortical  centres.      The  fornix  and  the  system 

of  the  ganglion  habenulae. 

The  fibres  of  the  stria  thalami  which  do  not  end  within  the  ganglion  habenulae 
traverse  the  latter  and  enter  the  commissura  interhabenularis — a  bundle  of  transverse  fibres 
lying  in  front  of  the  glandula  pinealis.  Some  of  these  fibres  end  in  the  ganglion  of  the 
opposite  side,  others  pass  to  the  roof  of  the  mid-brain,  especially  to  the  superior  colliculus, 
while  still  others,  perhaps,  come  into  relation  with  the  posterior  longitudinal  bundle. 

Within  the  ganglion  habenulae,  the  fasciculus  retroflexus  of  Meynert  takes  its 
origin.  This  bundle  ends  within  the  substantia  perforata  posterior,  in  the  region  imme- 
diately in  front  of  the  pons,  in  a  small  nucleus,  the  ganglion  interpedunculare  of  Gudden. 
The  bundle  is  called,  therefore,  also  the  tractus  habenulo-peduncularis. 

Within  the  ganglion  interpedunculare  arises  the  tegmental  tract  of  the  interpedun- 
cular  ganglion.  The  fibres  pass  dorsalward  as  far  as  the  central  gray  to  end  partly  in  the 
ganglion  tegmenti  profundum  and  partly  in  the  ganglion  tegmenti  dorsale  and  the  surround- 
ing central  gray  substance.  Here  joins,  in  turn,  the  dorsal  longitudinal  bundle  of  Schiitz. 


PATHS   OF  THE   RHINENCEPHALCN. 


149 


3.  Connection   of  the  primary   centres   of  the    two    sides. — The   fibres   arise 
within   the  cortex   of  the   tractus  olfactorius  and  pass,   forming   the  pars  olfactoria  of  the 
anterior  commissure,   to  the  tractus  of  the  opposite  side.      Here  they  end,   partly  within 
the  granule-layer  and  within  the  locality  of  the  olfactory  glomeruli  of  the  bulbus. 

4.  Further   connections    of  the    primary   centres. — Direct   fibres  pass   to  the 
tuber  cinereum,   to  the   corpus  mamillare,   to  the  lower   lying  brain -segments  and  to  the 
spinal    cord.      They  form  the   olfactory  radiation  to    the   diencephalon   and   to    the    mid- 
brain — tractus  olfacto-mesencephalicus,  the  basal  olfactory  bundle  of  Wallenberg. 

The  tract  of  fibres  passing  to  the  corpus  mamillare  is  further  joined  by  the  fibre- 
system  of  the  mammillary  body,  whereby  further  relations  with  the  thalamus  and  the 
mid-brain  are  established.  A  similar  connection  of  the  primary  centres  is  effected  through 
the  fibre-system  of  the  ganglion  habenulae. 


Fuse,  tegmento-mamillaris 
(Gtidden's  tegmental  strand) 


Fasc.  thalamo-mamillaris 
(  Vicg  d'Azyr's  bundle) 


Tract    olfacto-mesenceph. 
(  Wallenberg) 

Tuber  cinereu 

Fasc.  mamillaris  frinceps 


Corf,, 
mamllL 


141. — Fibre-tracts  of  the  rhinencephalon.      Further   connections  of  the  prir 
Wallenberg  and  the  system  of   the  corpus  mar 


Peduncnhis  corpor.  mamillaris 
Ganglion  in  terpedim  en  la  re 


ary  eentres.      Basal  olfactory  bundle  of 
illare. 


Ascending  fibres  from  the  lower  brain-segments,  as  strands  from  the  vicinity  of  the  end- 
nucleus  of  the  trigeminus,  are  also  credited  with  terminating  within  the  primary  olfactory 
centres.  Since  terminal  arborizations  of  the  trigeminus  are  found  within  the  regio  olfactoria 
of  the  nasal  mucous  membrane  and  since  this  nerve,  perhaps,  also  shares  in  the  conduction 
of  olfactory  stimuli,  it  is  not  impossible  that  impulses  may  be  carried  from  the  olfactory 
region  to  the  cortical  olfactory  centre  by  means  of  the  ascending  central  trigeminal  tract. 

5.  Connection   of  the    cortical    centres    of  the   two    sides. — This   is  accom- 
plished  by  the    fibres   of   the   fornix    transversus,   and,    perhaps,   by  the    pars    interhemi- 
sphaerica  of  the  anterior  commissure. 

6.  Further  connections  of  the   cortical  centres. — The  fornix  periphericus  of 
Arnold,  or  the  cingulum,  is  to  be  regarded  as  an  association  bundle  of  the  rhinencephalon. 
It  appears  as  an  arcuate  bundle,  which  surrounds  the  rostrum,   knee,  body  and  splenium 
of  the  corpus  callosum  ;   at  the  isthmus  it  becomes  narrow  and  expands  toward  the  front 
end  of  the  uncus.      It   consists  of  fibres  which  do  not  extend   the  entire   length    of  the 
tract,   but  form  larger  or   shorter  strands,   whose   crooked  ends  radiate  within  the  white 


i5o  THE   FIBRE-TRACTS. 

substance  of  the  neighboring  convolutions.  The  cingulum  appears  to  be,  therefore,  not 
properly  an  association  bundle  of  the  rhinencephalon,  but  an  association  strand  of  the 
different  convolutions  of  the  medial  surface  of  the  hemisphere  (Fig.  126). 

Reviewing  the  entire  fibre-tracts  of  the  rhinencephalon,  we  recognize,  in  the  first 
place,  that  a  centripetal  projection  path  conducts  the  impulse  from  the  regio  olfactoria  to 
the  primary  centres  and  thence  to  the  cortical  centre  proper  ;  secondly,  that  a  centrifugal 
projection  path  transfers  impulses  from  the  cortical  olfactory  centre  to  subcortical  centres 
(corpus  mamillare,  ganglion  habenulae),  from  which  latter  then,  by  means  of  further 
paths,  still  other  nuclei  may  be  influenced.  Thirdly,  the  tracts  that  arise  within  the 
primary  centres  and  pass  directly  to  the  subcortical  ganglia,  constitute  special  reflex  paths, 
and  by  means  of  these,  in  consequence  of  the  transference  of  the  impulse  to  the  most 
diverse  nuclei  of  the  brain-stem,  as  the  nuclei  of  the  motor  nerves,  the  most  varied 
reflex  movements  may  be  induced.  Finally,  the  peripheral  and  central  districts  of  the 
rhinencephalon  of  both  hemispheres  are  brought  into  relation  with  each  other  by  means 
of  certain  systems  of  commissural  fibres  ;  through  the  fornix  periphericus,  the  central 
district  is  also  connected  with  the  adjoining  regions  of  the  pallium. 

CONDUCTION   PATHS  OF  THE  DIENCEPHALON. 

The  connections  which  unite  the  diencephalon  with  other  parts  of  the  brain  have, 
in  large  part,  been  presented  in  the  preceding  section.  To  these  belong,  in  the  first 
place,  the  tractus  cortico-thalamici  and  thalamo-corticales — the  thalamic  peduncles — of  which 
the  tegmental  tract  and  the  optic  radiation  may  be  again  mentioned  as  of  especial 
importance.  Others  to  be  recalled  are  those  fibre-tracts  which  unite  certain  parts  of  the 
olfactory  brain  with  the  thalamencephalon  and  the  hypothalamus — fornix,  stria  medullaris, 
basal  olfactory  bundle,  which  tracts  are  joined,  moreover,  by  those  passing  from  the 
diencephalon  to  the  mid-brain — fasciculus  mamillo-tegmentalis,  pedunculus  corporis  mamil- 
laris,  tractus  habenulo-peduncularis.  Further,  the  connections  which  join  the  corpus 
striatum  with  the  thalamus  and  with  the  subthalamic  ^region — the  radiatio  strio-thalamica 
and  strio-subthalamica. 

Within  the  pulvinar  thalami  and  the  corpus  geniculatum  laterale,  which  parts, 
together  with  the  superior  colliculus,  constitute  the  primary  visual  centre  (page  17?),  the 
fibres  of  the  tractus  opticus  end.  The  corpus  geniculatum  mediale,  with  the  inferior 
colliculus,  constitutes  the  primary  auditory  centre,  since  within  these  parts,  particularly 
within  the  medial  geniculate  body,  the  fibres  of  the  lateral  fillet  end  ;  the  latter,  as  later 
to  be  described  (page  179),  serves  to  conduct  the  impulses  from  the  end-nuclei  of  the 
acoustic  nerve  farther  centrally  and,  hence,  represents  the  primary  auditory  tract.  From 
the  corpus  geniculatum  laterale  and  the  pulvinar  thalami,  the  optic  radiation  passes  to 
the  cortex  of  the  visual  centre  in  the  occipital  lobe  ;  from  the  corpus  geniculatum  mediale, 
the  secondary  auditory  tract  passes  to  the  cortex  of  the  auditory  centre  in  the  temporal  lobe. 

Within  the  thalamus,  moreover,  end  certain  fibre-strands  which  corne  from  the 
cerebellum,  the  medulla  oblongata  and  the  spinal  cord.  Bundles  of  fibres,  from  the 
nucleus  dentatus  and  in  small  part  also  from  the  nucleus  tecti,  pass  forward  from  the 
cerebellum,  constituting  collectively  the  superior  cerebellar  peduncle.  The  larger  part  of 
these  fibres,  after  decussation,  reaches  the  nucleus  ruber  in  the  tegmentum  of  the  mid- 
brain  and  there  ends — tracius  cerebello-tegmentalis ;  the  smaller  part  of  the  fibres  passes 


PATHS   OF  THE   MESENCEPHALON.  151 

directly  to  the  thalamus,  joining  such  as  come  from  the  red  nucleus — tractus  rubro- 
thalamicus.  The  fibres  proceeding  from  the  medulla  oblongata  and  from  the  spinal  cord 
form  the  large  ascending  sensory  path,  tractus  spino-  et  bulbo-thalamicus ,  the  detailed 
origin  and  course  of  which  will  be  later  considered.  For  the  present  suffice  it  to  note, 
that  this  tract,  known  as  the  medial  fillet  or  lemniscus  media/is,  carries  fibres  from  the 
spinal  cord,  the  nuclei  of  the  posterior  column  and  the  end-nuclei  of  the  sensory 
cerebral  nerves.  Its  termination  is  principally  within  the  lateral  nucleus  and  the  centrum 
medianum  of  the  thalamus.  Impulses  from  the  spinal  cord,  the  nuclei  of  the  posterior 
column  and  the  olivary  nucleus  of  the  medulla  may  reach  the  thalamus  also  by  way  of 
the  cerebellum  and  the  superior  cerebellar  peduncle.  These  paths  are  shown  in  Figs. 
142  and  147.  The  tractus  thalamo-olivaris  is  a  spinalward  coursing  path,  that  connects 
the  thalamus  with  the  olive  of  the  medulla  (Fig.  147);  it  is  also  termed  the  central 
tegmental  tract.  Since  the  olive  sends  fibres  to  the  cerebellum,  impulses  from  the 
thalamus  may  be  conveyed  to  the  cerebellum  by  this  tract. 

Probably  still  other  paths  proceed  from  the  thalamus  downward,  to  end  within  the  mid- 
brain,  the  pons,  the  medulla  and  the  spinal  cord.  Such  connections,  however,  are  not  accu- 
rately determined.  The  descending  tractus  thalamo-spinalis  accompanies  the  tractus  rubro- 
spinalis  to  the  spinal  cord,  within  which  it  courses  in  the  dorsal  part  of  the  lateral  column. 

Finally,  mention  must  be  made  of  the  system  of  the  ventricular  gray,  which  gray 
substance  covers  the  medial  surface  of  the  thalamus  and  hypothalamus  and  the  floor  of 
the  third  ventricle  and  is  continuous  with  the  gray  substance  surrounding  the  aquaeductus 
Sylvii  and  investing  the  floor  of  the  fourth  ventricle.  The  fibres  from  the  cells  situated 
within  this  gray  substance  pass  to  all  the  thalamic  nuclei,  while  delicate  longitudinal 
strands  proceed  caudalward  through  the  gray  in  the  medulla  oblongata  and  into  the 
spinal  cord.  This  system  of  longitudinal  fibres,  which  has  been  already  noticed  in 
connection  with  the  fibre-tracts  of  the  rhinencephalon,  is  the  fasciculus  longitudinalis 
dorsalis,  or  dorsal  longitudinal  bundle  of  Schiitz,  and  is  closely  connected  with  the 
nuclei  of  the  cerebral  nerves  and  other  ganglia.  Concerning  the  significance  of  the 
entire  system,  we  are,  at  present,  insufficiently  informed.  According  to  Edinger,  it  is 
not  unlikely,  that  all  these  nuclei  and  fibres  constitute  a  central  apparatus  of  the  sympathetic. 

CONDUCTION    PATHS  OF  THE  MESENCEPHALON. 

The  mesencephalon,  which,  as  the  smallest  of  the  brain  segments,  includes  the 
quadrigeminal  region  and  the  cerebral  peduncles,  is  traversed  by  several  main  tracts,  on 
the  one  hand,  and  is  the  termination  or  the  origin  of  many  fibre-strands,  on  the  other. 

I.  The  chief  tracts  traversing  the  mid-brain  are  those  which  descend  from 
the  cortex  of  the  cerebral  hemispheres,  already  described  in  connection  with  the  conduc- 
tion paths  of  the  telencephalon,  namely :  the  frontal  pantile  tract,  the  occipito-lemporal 
pontile  tract  and  the  motor  tract.  These  three  chief  paths  pass  through  the  basis  pedun- 
culi  or  crusta,  the  frontal  fibres  occupying  the  medial  and  the  occipito-temporal  the 
lateral  part,  while  the  motor  tract  appropriates  the  middle  portion  of  the  crusta  between  the 
pontile  tracts.  An  additional  traversing  path  is  the  sensory  tract  or  medial  fillet,  which 
ascends  from  the  spinal  cord,  the  nuclei  of  the  posterior  columns  and  the  end-nuclei  of  the 
sensory  cerebral  nerves  and  continues  to  the  thalamus,  joined  by  the  tegmental  tract  that 
unites  the  thalamus  with  the  sensory  region,  in  the  cortex  of  the  parietal  lobe.  This 


152 


THE   FIBRE-TRACTS. 


ascending  sensory  tract,  however,  does  not  pass  through  the  crusta  of  the  cerebral  pe- 
duncle, but  through  the  tegmental  area.  Of  the  additional  traversing  paths,  the  tractus 
thalamo-olivaris  or  the  central  tegmental  tract  deserves  special  mention. 


Tract,  cerebro-spinalls 
Tractus  tecto-spinalis  and  spino-tectalis 


FIG    142. — Schematic  representation   of  the  chief  connections   of  the   mid-brain   and  of  the  tracts  passing 
•  through  the   mid-brain. 

II.     Tracts  ending  within  the  mid-brain: 

a.  Within  the  region  of  the  superior  colliculus  end  some  fibres  of  the  optic  tract ; 
within  the  inferior  colliculus,  the  fibres  or  collaterals  of  the  lateral  fillet,  which  latter 
represents  the  primary  auditory  path. 


PATHS   OF  THE   MESENCEPHALON.  153 

b.  Within  the  quadrigeminal  region   end  additional    fibres  from   the  cortex,   tractus 
cortico-tectales — in  the  superior   colliculus  principally    fibres,  from  the  occipital  lobe  and  in 
the  inferior  colliculus  those  from  the  temporal  lobe  ;  further  fibres  are  those  of  the  traclus 
spino-tectales,  which  ascend  from  the  lateral  column  of  the  spinal  cord. 

c.  Within  the  nucleus  ruber  end,  first,  fibres  from  the  cortex  (frontal  lobe,  regio  opercu- 
laris)  and  from  the  corpus  striatum;  second  and  most  important,  fibres  from  the  cereDellum. 
The  last  take  their  origin  in  the  nucleus  dentatus,  in  small  part  also  in  the  nucleus  tecti,  and, 
perhaps,  also  in  the  cortex  of  the  cerebellum,  and  form  the  brachia  conjunctiva  or  superior 
cerebellar  peduncles.     After  decussation  within  the  tegmentum  of  the  mid-brain,  the  fibres 
end  in  the   nucleus  ruber  and,  in  part,  also  in  the  thalamus — tractus  cerebello-tegmentalis. 

d.  Small  individual  fibre-bundles,   which  in  part  end  in  the  mid-brain  and  in  part 
run  still  farther  caudally — the  tegmental    bundle  of    the    ganglion  interpedunculare,   fibre- 
strands  from  the  corpus  mamillare  and  from  the  posterior  longitudinal  bundle. 

III.  Tracts  arising  within  the  mid-brain: 

a.  Tractus   tecto-bulbaris   et  tecto-spinalis,    fibre-strands    which    arise   frorn  the  deep 
medullary  substance  of  the  corpora  quadrigemina,  cross  (Meynert's  tegmental  decussation) 
and    terminate  within  the  nuclei  of   the  medulla    oblorigata  and    within    the   anterior  and 
lateral    columns    of    the    spinal    cord.       Since   fibres    of  the   visual   tract   end   within   the 
superior    colliculus    and    those   of    the    auditory    tract    within    the    inferior   colliculus,   the 
impulses  brought  to  the  mid-brain  by   these   paths  may  be  conveyed  to  the  medulla  and 
the  cord  by  the    tecto-bulbar  and  the  tecto-spinal  tract  respectively.     These  paths,  there- 
fore, are  also  called    the    visuo-acoustic  reflex  tract.     The    tract   leading    to    the   anterior 
column  is  known  also  as  the  fasciculus  longitudinalis  praedorsalis,  since   the  bundle  lies 
ventral  to  the  posterior  longitudinal  bundle  in  its  course  through  the  brain-stem. 

b.  Tractus  tecto-cerebellarcs,  from  the   quadrigeminal  plate  to  the  cerebellum. 

c.  Tractus    tecto-pontinus    (Munzer},    a   small    fibre-strand    that    arises    within    the 
quadrigeminal    region,    more    especially  within    the    inferior    colliculus,    and    ends    in    the 
pontile  nuclei  in  the  vicinity  of  the  pyramidal  tract. 

A  small  bundle,  the  tractus  tecto-reticularis  of  Pavlow,  extends  from  the  quadrigem- 
inal region  to  the  tegmentum  of  the  pons  and  ends  within  the  nucleus  reticularis  tegmenti. 

d.  Tractus    rubro-spinalis,    also    known    as    Monakow" s    bundle,  arises    in    the   red 
nucleus  of   the  tegmentum.      The  fibres    emerging  from    the    nucleus    cross   and    descend 
through  the  pontile    tegmentum  and  the  medulla    oblongata  to  the  lateral   column  of   the 
spinal  cord,   to  end  within  the  anterior  horn. 

Moreover,  the  tractus  rubro-reticularis  includes  fibres  which  pass,  crossed  and  un- 
crossed, from  the  red  nucleus  to  the  formatio  reticularis  of  the  pons  and  of  the  medulla 
oblongata  ;  a  further  bundle,  the  tractus  rubro-laquearis,  passes  from  the  red  nucleus  to 
the  nucleus  of  the  lateral  fillet. 

c.  Fasciculus  longitudinalis  medialis,  commonly  called  the  posterior  longitudinal 
bundle,  is  composed  of  fibres  taking  their  origin  in  different  places.  The  principal  fibre- 
strands  arise  from  Deiters'  nucleus  and  from  the  nucleus  of  the  posterior  commissure  and 
of  the  posterior  longitudinal  bundle,  situated  in  front  of  the  oculomotor  nucleus  (p.  182). 

f.  Finally  it  must  be  noted,  that  the  oculomotor  and  the  trochlear  nerves,  as  well 
as  a  small  motor  root  of  the  trigeminus,  have  their  origin  within  the  mid-brain. 


THE   FIBRE-TRACTS. 


CONDUCTION  PATHS  OF  THE  METENCEPHALON. 

Before  considering  the  individual  fibre-tracts  connecting  the  cerebellum  and  the  pons 
with  other  parts  of  the  brain  and  the  spinal  cord,  the  structure  of  the  cerebellar  cortex 
must  be  more  closely  examined. 

HISTOLOGY   OF   THE  CEREBELLAR   CORTEX. 

The  cerebellar  cortex  presents  the  following  layers  : 

1.  The  molecular  /ayer—the  outermost  stratum  ; 

2.  The  layer  of  Purkinje  cells — the  middle  stratum  ; 

3.  The  granule  layer — the  innermost  stratum. 


Purkinje  cells 


Basket  cellf  -  - 


Moss  fibres  •. 


Sagittal  section 
PIG.  143. — Schematic  representation  of  the  cerebellar  cortex. 

The  Purkinje  cells  send  their  richly  branched  protoplasmic  processes  or  dendrites 
into  the  molecular  layer,  while  the  axones  of  the  cells  pass  through  the  granule  layer  to 
the  white  substance  of  the  cerebellum. 

Within  the  molecular  layer,  in  addition  to  small  cortical  cells  with  short 
axones,  are  found  the  basket-cells.  The  latter  are  distinguished  by  their  axones, 
which  run  sagittally  and  parallel  to  the  surface  and  give  off  numerous  collaterals 
that  pass  inward  and  surround  the  cell-bodies  of  the  Purkinje  cells  with  basket-like 
ramifications  (Fig.  143). 

Within  the  granule  layer,  the  small  granule-cells  are  the  chief  elements.  They  are 
small  spherical  cells  with  from  three  to  five  short  dendrites.  Their  axones  pass  into  the  molec- 


PATHS   OF  THE   CEREBELLUM. 


155 


.ular  layer,  where  they  divide  into  two  branches,  that  run  parallel  to  the  surface  and  in  corre- 
spondence with  the  direction  of  the  cerebellar  convolutions  or  folia.  These  axones 
extend,  therefore,  in  the  frontal  plane  and  not,  as  do  those  of  the  basket-cells,  in  the 
sagittal  plane.  During  their  course,  the  branches  give  off  collaterals  which  pass  to  the 
Purkinje  cells.  In  addition  to  the  granule-cells,  Golgi  II  type  cells  occur,  whose 
dendrites  often  extend  far  into  the  molecular  layer  and  whose  axones  resolve  into 
branchings  of  unusual  richness. 

Nerve-fibres  enter  the  cortex  from  the  subjacent  white  substance.  Of  these,  the 
' '  climbing  fibres' '  pass  to  the  molecular  layer  and  there  end  among  the  dendrites  of  the 
Purkinje-cells,  while  the  "moss  fibres"  terminate  chiefly  within  the  granule  layer.  Im- 
pulses, conveyed  by  these  fibres  which  enter  the  cerebellum,  are  transferred  to  the 
different  varieties  of  cortical  cells.  In  this  connection,  it  is  worthy  of  special  note,  that 
by  means  of  the  basket-cells  impulses  are  conveyed  in  the  sagittal  direction  and  by  means 
of  the  granule-cells  in  the  frontal  direction  and  transferred  to  numerous  Purkinje  cells. 


FIBRE-TRACTS   OF   THE   CEREBELLUM. 

All  cortical  regions  of  the  cerebellum  are  linked  together  by  means  of  arched 
fibres,  the  fibrae  arciformes.  Such  association  systems  unite  neighboring  folia  or  lobules 
of  the  cerebellum.  The  cortex,  moreover,  sends  centrifugal  fibres  to  the  nuclei — to  the 
nucleus  dentatus  and  the  nucleus  fastigii,  as  well  as  to  Deiters'  nucleus. 

The   chief  connections  of  the  cerebellum  are: 

i,   Tractus  ponto-cerebel lares,    composed    of   fibres    which  arise    in  the  pontile  nuclei 
and  proceed    to    the    cerebellar    hemisphere  of  the    opposite  side.     These  tracts  form  the 
middle  cerebellar  peduncle.     Since  the  ponto-cerebellar  tract  continues  the  tractus  corticis 
ad  pontem,  connecting  the  cere- 
bral    cortex     with     the     pontile 
nucleus,     impulses     are     carried 
from  the  cerebrum   to    the   cere- 
bellum.    Relations    between    the 
cerebral    cortex    and    the     cere- 
bellum may  be  further  established 
by    way    of     the    thalamus    and 
the    inferior  olive. 

Some  fibres  pass  in  the 
opposite  direction,  from  the  cere- 
bellum through  the  middle  cere- 
bellar peduncle  to  the  pons  and 


Thalamus 


Nucleus  ritber 


Tractus  cerebello-tegmentales 
(Superior  peduncle) 


from  -which  pass  the  ponto-cerebellar  tracts 

FIG.  144. — Fibre-tracts  of  the  cerebellum. 


thence,  as  the  fibrae  rectae  pontis, 
dorsally  within  the  raphe  of  the 
pons  to  the  centro -lateral  nucleus 

recticularis    tegmenti  pontis,    thus    constituting    the    tractus    cerebello-tegmentalis   pontis. 

2.   Tractus   cerebello-tegmentales,    composed   of    fibres    which  arise    in    the    nucleus 

dentatus  and  partly  in  the  nucleus  fastigii    or    roof-nucleus    of   the    cerebellum,  pass    for- 


156 


THE   FIBRE-TRACTS. 


ward,  decussate  in  the  quadrigeminal  region  and  .end  within  the  nucleus  ruber  or  the 
thalamus.  They  constitute  the  superior  cerebellar  peduncle  or  brachium  conjunctiva, 
the  crossing  being  known  as  the  decussation  of  the  superior  peduncle.  These  fibre- 
bundles,  chiefly  from  the  nucleus  dentatus  cerebelli,  give  off  descending  collateral 
branches,  which  may  be  followed  as  a  special  bundle  as  far  as  the  pons  and  the 
medulla  oblongata,  where  they  probably  end  in  motor  nuclei.  In  addition  to  this 


Jrttctits  corticis 
ad  fontem 


FIG    145. — Connections  of  the  cerebral  cortex  with  the  cerebellum  and  of  the  cerebellum  with  the  cerebral  cortex. 


robust  cerebellofugal  tract,  other  efferent  bundles  pass  caudalward  from  the  roof- 
nucleus,  of  the  same  and  the  opposite  side,  into  the  tegmental  area  of  the  medulla 
oblongata,  where  they  end  around  the  cells  of  the  substantia  reticularis.  These  fibres, 
however,  emerge  from  the  cerebellum  by  way  of  the  inferior  cerebellar  peduncle  or 
restiform  body  and  are  specially  designated  as  the  tractus  cerebello-tegmenlalis  bulbi, 
while  those  in  the  superior  peduncle  are  called  the  tractus  cerebello-tegmentalis 
mesencephali. 


PATHS    OF  THE   CEREBELLUM. 


157 


By  means  of  the  middle  cerebellar  peduncles,  therefore,  especially  impulses  from 
the  cerebrum  are  conveyed  to  the  cerebellum.  The  superior  cerebellar  peduncle,  .on 
the  contrary,  carries  impulses  by  way  of  the  red  nucleus  and  the  thalamus  from  the 
cerebellum  to  the  cerebrum.  In  addition,  however,  by  way  of  the  tractus  cerebello- 
tegmentalis  mesencephali  (cerebellum  to  red  nucleus),  as  well  as  by  the  tractus  cerebello- 
tegmentalis  pontis  et  bulbi,  the  possibility  exists,  that  impulses  from  the  cerebellum 


libra  reticularis 


PIG,  146. — Tractus  cerebello-tegmentalis  mesencephali — -nucleus  dentatus  cerebelli — nucleus  ruber.  Tractus  cerebello- 
tegmentalis  pontis  et  bulbi.  Tractus  rubro-reticularis  and  tractus  rubro-spinalis ;  also  the  tracts  coursing  within  the 
formatio  reticularis. 

may  be  finally  transferred  to  the  motor  nuclei  of  the  cerebral  nerves  and  to  the 
gray  substance  of  the  spinal  cord.  Since  the  tractus  rubro-reticularis  passes  from 
the  red  nucleus  to  the  cells  of  the  formatio  reticularis  of  the  pons  and  of  the  medulla 
oblongata  and  the  tractus  rubro-spinalis  passes  to  the  spinal  cord,  while  other  fibres 
spring  from  the  cells  of  the  formatio  reticularis,  around  which  also  the  tractus  cere- 
bello-tegmentalis pontis  et  bulbi  end,  certain  fibre-strands  reach  far  down  into  the 
medulla  and  the  spinal  cord. 


158 


THE   FIBRE-TRACTS. 


3.  Constituents  of  the  inferior  cerebellar  peduncle,  fibres  which  come  from 
the  spinal  cord  and  the  medulla  oblongata  and  pass  to  the  cerebellum  by  way  of 
the  peduncle  or  restiform  body.  The  constitution  and  destination  of  the  latter 


Tract,  ceri'bello- 
tegmentalis 


Oliva 

Tractns  spino-olivari*  (Helwef) -\-A 

Tract,  splno-cerebiilaris  veiitritlis  (Gouoers)    .. 


Tractns  rubro-spinalis  (  Monakow ) 
Nuclei  fimic.  post. 
Tract,  fnnic  post 


Tract    splno  cerebellar  is  dor  sails 
( Flechsig) 


FIG.  147 — Schematic  representation  of  the  chief  connections  of  the  cerebellum. 

will     be     further     considered     in     connection      with      the     fibre-tracts     of     the     medulla 
oblongata    (page  170).  .    • 

The  chief  connections  of  the  pons  and  of  the  cerebellum  are  represented  in  Figs. 
144,  145  and  147.  It  is  to  be  noted,  further,  that  numerous  tracts,  descending  as  well 
as  ascending,  traverse  the  pons  (Figs.  142  and  147). 


CELL-GROUPS   OF  SPINAL  CORD. 


159 


THE  SPINAL  CORD. 

THE    GRAY    SUBSTANCE. 

Exclusive  of  the  supporting  tissue,  the  gray  substance  consists  principally  of  nerve- 
cells  with  their  protoplasmic  and  nerve-processes  and  nerve-fibres  ending  around  the 
nerve-cells.  Topographically  regarded,  four  different  cell-groups  may  be  distinguished. 
Thus,  in  the  anterior  horn  in  the  cervical  and  lumbar  enlargements,  a  ventro-medial 
and  a  ventro- lateral  and  a  dorso-medial  and  a  dorso-lateral  group  are  clearly  recognized  ; 
between  these  groups  lies  the  intermediate  zone  or  central  field  that  borders  the 
posterior  horn. 


Nerve-processes  from  tract-cells 


Spinal  ganglion 


Anterior  root 


Nerve-processes 
from  association  cells 


FIG.  148. — Schematic  representation  of  the  different  classes  of  cells  of  the  spinal  cord. 

Dorsal  to  the  dorso-.lateral  cells,  is  situated  the  cell-group  of  the  lateral  horn, 
while  Clarke's  column  lies  somewhat  medial  to  the  transition  of  the  intermediate  zone 
into  the  posterior  horn. 

Everywhere  within  the  posterior  horn,  mostly  small  cells  are  scattered  without 
the  definite  disposition  of  the  anterior  cornu ;  nevertheless,  also  here  different  groups 
are  defined,  as  the  basal,  the  central  and  the  marginal  cells  and  those  of  the 
substantia  Rolandi. 

In  contrast  to  this  subdivision  of  the  cord-cells  according  to  position  and  arrange- 
ment within  the  gray  substance,  a  classification  according  to  the  behavior  of  their  nerve- 


160  THE   FIBRE-TRACTS. 

processes  is  more  appropriate  for  a  presentation  of  the  fibre-tracts.     Therefore,  we  divide 
the  cells  of  the  spinal  cord  into  : 

1.  Cells,  whose    axones  pass  from   the   spinal  cord.      They  lie  within    the  anterior 
horn    and    are    called    the    motor   anterior    horn-cells.       Their    nerve-processes    form    the 
anterior  roots  emerging  from  the  spinal  cord. 

2.  Cells,     whose    axones  pass    into    the    white    substance.      Within    the    latter,    the 
axone    divides    into    an    ascending    and    a    descending    branch.     The   descending  branch, 
after    a    short    course,   again    enters    the    gray    substance,  where    it    ends;    the  ascending 
branch    courses  upward  within  the  white  substance.     These  cells  are  termed  column-cells, 
of  which  two  varieties  are  distinguished : 

a.  Cells,   whose  axones,   that    is,   ascending    branches,    pass    upward    and   as    special 
paths  connect  the  spinal  cord  with  the  brain.;  such  are  tract-cells. 

b.  Cells,    whose    ascending    branches    or    axones    after    a    longer    or    shorter  course 
again  enter  the  gray  substance  of  the  cord   and    serve    to  unite    different    cord-segments; 
such  are  association- cells. 

The  column-cells  may  be  further  distinguished  as  homolateral  and  contralateral. 
The  axones  of  the  former  pass  into  the  white  substance  of  the  same  side,  those  of  the 
latter  to  the  white  substance  of  the  opposite  side  by  way  of  the  anterior  commissure — 
commissure -cells.  The  column-cells  are  designated  respectively  as  anterior,  lateral  or 
posterior  column-cells  according  to  the  column  in  which  they  course.  While  the  column- 
cells  are  found  in  all  parts  of  the  gray  substance,  the  contralateral  column-cells  occur 
chiefly  within  the  base  of  the  posterior  horn  and  the  intermediate  zone.  (Commissure- 
cells  are  shown  in  the  first  and  second  cross-sections  in  Fig.  148.) 

3.  Cells  of  Golgi  II  type.       These   are  found  predominatingly  within  the    posterior 
horns  and  the  substantia  gelatinosa  Rolandi. 

The  motor  anterior  horn-cells,  whose  axones  form  the  motor  anterior  roots,  occupy 
a  special  position,  since  they  are  the  only  elements  that  send  their  axones  from  the  cen- 
tral organ  to  the  periphery.  The  column-cells  and  the  Golgi  cells,  with  their  entire 
expansions,  belong  to  the  central  nervous  system.  The  tract-cells  establish  relations  between 
the  spinal  cord  and  the  higher  lying  centres  ;  the  association-cells  serve  to  transfer  an 
impulse  received  within  the  cord  to  higher  and  lower  lying  cell-complexes  ;  while  the  field 
of  activity  of  the  cells  of  Golgi' s  II  type  is  limited  to  the  immediate  vicinity.  The  nerve- 
processes  of  the  association-cells  are  also  spoken  of  as  endogenous  fibres. 

THE  WHITE  SUBSTANCE. 

The  white  substance  consists  essentially  of  the  longitudinally  coursing  nerve-fibres.. 
The  following  chief  systems  of  fibres  are  distinguished  : 

a.  Fibres,  which   arise   within    the   cerebral  cortex   and  within    certain  parts  of   the 
brain,  descend  within  the  spinal  cord  and  there  end. 

b.  Fibres,  which    arise  within    the   gray  substance    of   the    spinal    cord   and    end    in 
higher  lying  parts — axones  of  the  tract-cells. 

c.  Fibres,  which  unite  particular  levels  of  the  spinal  cord — axones  of  the  association-cells. 

d.  Fibres,  which   arise  from    the   spinal   ganglia   as  continuations    of    the    posterior 
roots,  enter  the  spinal  cord  and  course  within  the  posterior  columns. 


PATHS   OF  THE  SPINAL   CORD.  161 


1.  TRACTS  OF  THE  ANTERIOR  COLUMN. 

The  tractus  cerebro-spinalis  anterior  or  the  anterior  pyramidal  tract  passes  medially, 
along  the  anterior  median  fissure.  The  fibres  end,  after  crossing  in  the  anterior  commis- 
sure, within  the  anterior  horn  of  the  opposite  side. 

The  field  of  the  anterior  pyramidal  tract  is  shared  by  fibres  that  descend  from  the 
mid-brain.  They  constitute  the  tractus  tecto-spinalis,  or  fasciculus  sulco-marginalis.  The 
fibres,  in  part,  cross  after  their  origin  in  the  corpora  quadrigemina.  A  portion  of  the 
fibres  pass  also  to  the  lateral  column  of  the  cord — tractus  tecto-spinalis  lateralis. 

Burdach's       Coil's         Oval 
Root-zone       strand          strand     bundle     Ventral  field 

Comma  bundle 

teral  pyramidal  tract 

Tractus  spino-cerebellaris  dor- 
salts  (Flechsig) 

Tractus  spino-cerebellaris  ven- 
tralis  (Cowers) 

-Tractus  spino-olivaris  (Helweg) 

Anterior  ground  bundle 

Fasciculus  Tract,  vestibulo-spinalis 

sulco-marginalis 
Anterior  pyramidal  tract    s.  Tract,  tecto-spinalis 

FIG.  149. — Fibre-systems  of  the  white  substance  of  the  spinal  cord 

At  the  ventral  border  of  the  anterior  column,  fibres  are  found  which  come  from 
Deiters'  nucleus  and  constitute  the  tractus  vestibulo-spinalis  anterior  or  the  anterior  mar- 
ginal bundle. 

Lastly,  as  an  additional  descending  path  within  the  anterior  column,  the  posterior 
longitudinal  bundle,  or  the  fasciculus  longitudinalis  medialis,  must  be  noted. 

The  remaining  part  of  the  anterior  column  constitutes  the  anterior  ground  bundle 
or  fasciculus  anterior  propriiis.  Within  this  field  longitudinally  course  the  endogenous 
fibres,  which  serve  to  unite  different  segments  of  the  spinal  cord. 

2.  TRACTS  OF  THE  LATERAL  COLUMN. 

The  tractus  cerebro-spinalis  lateralis  or  the  lateral  pyramidal  tract  extends  as  a 
robust  bundle  in  the  dorsal  part  of  the  lateral  column.  The  termination  of  the  fibres  is 
in  the  anterior  horn  of  the  same  side  of  the  cord. 

The  tractus  spino-cerebellaris  dorsalis  or  direct  cerebellar  tract  lies  at  the  periphery, 
lateral  to  the  pyramidal  tract.  The  fibres  arise  from  the  cells  of  Clarke's  column,  extend 
upward  within  the  lateral  column  and  pass,  as  a  constituent  of  the  restiform  body,  to  the 
cerebellum,  where  they  end  within  the  anterior  superior  worm. 

The  tractus  spino-cerebellaris  ventralis  or  Cowers*  tract  also  lies  at  the  periphery 
of  the  lateral  column,  in  front  of  the  direct  cerebellar  tract,  and  likewise  ends  within  the 


•62 


THE   FIBRE-TRACTS. 


cerebellum.  The  fibres  take  their  origin  from  cells,  on  the  same  and  the  opposite  side, 
which  lie  in  the  lateral  part  of  the  posterior  horn  and  in  the  central  field  of  the  gray 
substance.  They  ascend  at  first  with  the  direct  cerebellar  tract,  do  not,  however,  enter 
the  restiform  body,  but  continue  as  far  as  the  pons,  then  enter  the  superior  cerebellar 
peduncle  and  pass  backward  to  the  cerebellum,  where  they  end  within  the  anterior  part 
of  the  superior  worm  (Fig.  74,  fibrae  arciformes). 

The  tractus  spirw-olivaris  or  triangular  bundle  of  Helweg  is  an  additional  small 
tract,  which  lies  at  the  periphery  of  the  lateral  column,  ventral  to  Gowers'  bundle,  and 
establishes  relations  between  the  spinal  cord  and  the  inferior  olivary  nucleus  in  the 


Fibres  of 
lateral  bundl 


FIG.  150. — Diagram  of  the  tracts  of  the  posterior  column. 

medulla  oblongata.  The  source  of  the  fibres  has  not  been  positively  determined.  They 
form,  perhaps,  an  ascending  tract,  arising  within  the  gray  substance  of  the  cord  and 
ending  in  the  olive;  according  to  other  views,  the  bundle  conveys  both  ascending  and 
descending  fibres.  The  entire  area  of  the  lateral  column  included  between  the  foregoing 
tracts  and  the  gray  substance,  belongs  to  the  lateral  ground  bundle  or  fasciculus  lateralis 
proprius.  Within  this  again  appear  numerous  endogenous  fibres,  long  and  short  associ- 
ation fibres,  which  bind  together  various  higher  and  lower  lying  segments  of  the  spinal 
cord.  The  short  fibres  lie  close  to  the  gray  substance  and  form  the  lateral  boundary 
zone.  In  addition,  within  the  lateral  ground  bundle  are  found  the  following  sets  of  fibres: 
the  tractus  rubro-spinalis  or  Monakow's  bundle,  whose  fibres  descend  from  the  nucleus 
ruber  of  the  opposite  side  and  lie  within  the  cord  medial  to  the  direct  cerebellar  and 
ventral  to  the  lateral  pyramidal  tract,  partly  within  the  field  of  the  latter.  The  tractus 


PATHS   OF  THE   SPINAL   CORD. 


163 


Gall's  coin* 


Burdach's  column 


vestibtdo-spinalis  lateralis,  from  Deiters'  nucleus,  passes  somewhat  more  ventral.  Medial 
to  Cowers'  bundle  lies  the  tractus  spino-thalamicus .  The  fibres  of  this  path  are  the 
axones  of  the  commissure-cells  of  the  cord,  which  pass  through  the  anterior  commissure 
to  the  opposite  lateral  column  and  there  turn  upward.  The  termination  of  the  path  is 
in  the  thalamus.  The  tractus  spino-tectalis  consists  of  a  fibre-strand,  that  accompanies  the 
spino-thalamic  tract  and  ends  in  the  region  of  the  corpora  quadrigemina.  The  entire 
path  is,  therefore,  also  called  the  tractus  spino-tectalis  et  thalamicus.  Approximately  within 
the  same  field,  the  tractus  tecto-spinalis  lateralis  descends  from  the  quadrigeminal  region; 
likewise,  in  the  vicinity  of  the  spino-thalamic  fibres,  the  tractus  thalamo-spinalis  descends 
as  a  path  from  the  thalamus. 

3.   TRACTS  OF  THE   POSTERIOR  COLUMN. 

The  posterior  column  is  composed,  in  largest  part,  of  the  continuations  of  the 
sensory  posterior  root-fibres,  that  proceed  from  the  spinal  ganglia  (Fig.  150).  The  cells 
of  the  latter  give  off  a  nerve-process,  which  soon 
divides  into  two  branches,  one  passing  peripheral- 
ward  and  the  other  centralward.  The  centrally 
coursing  branches  enter  the  spinal  cord,  as  the  pos- 
terior root,  as  two  more  or  less  distinct  bundles. 
One  of  these  is  made  up  of  fine  fibres,  lies  lateral 
and  passes  toward  the  substantia  gelatinosa  Ro- 
landi;  the  other  consists  of  coarse  fibres,  lies  medial 
and  passes-  toward  the  posterior  column.  The  en- 
trance zone  of  the  lateral  bundle,  between  the  apex 
of  the  posterior  horn  and  the  periphery  of  the  cord, 
is  known  as  Lissauer's  marginal  zone;  that  of  the 
medial  bundle,  to  the  inner  side  of  the  posterior 
horn,  is  called  the  radicular  zone.  Immediately 
upon  entering  the  spinal  cord,  the  fibres  of  both 
bundles  undergo  a  Y-form  branching.  Both  result- 
ing branches  assume  a  longitudinal  direction  and 
during  their  course,  respectively  up  or  down,  give 
off  numerous  collaterals  to  the  gray  substance  of  the 
cord.  The  descending  branch  is  the  thinner  and 
ends,  after  a  short  course,  within  the  gray  substance. 
According  to  their  length,  the  ascending  fibres  are 
short,  medium  or  long.  The  short  fibres  pass  into 
the  gray  substance  after  a  very  limited  course;  those 
of  medium  length  proceed  farther  upward,  but  like- 
wise end  within  the  cord  by  bending  over  into  the 
gray  substance;  while  the  long  fibres  ascend  to  the 
medulla  oblongata,  where  they  end  within  the  poste- 
rior column  nuclei,  the  nucleus  gracilis  and  cuneatus. 

The   fibres   entering   the   cord   below  are   dis-  FIG.  151.— Diagram  illustrating  the  gradual 

i          j  i  11  •  i   «•  medial  displacement    of   the   long   tracts    of   the 

placed  more  and  more  towards  the  mid-line  by  the      posterior  column. 


i64 


THE   FIBRE-TRACTS. 


fibres  entering  at  higher  levels  ;  those  fibres,  therefore,  that  on  entering  the  cord  occupy  the 
lateral  part  of  the  posterior  column, as  they  ascend  soon  collectively  constitute  the  middle  and, 
finally,  the  innermost  part  of  the  column.  Hence,  as  already  .noted,  the  posterior  column 
exhibits  in  the  cervical  region  of  the  cord,  a  subdivision  into  the  medial  fasciculus  gracilis 


LJ  Nucl.ruber 

Corp.  qua  drigem 


Tract,  vestibulo-spinalis  anterior 


. .  Tract,  vestibulo-spinalis  laterali. 

\ 

\ Tract,  rulro-spinalis 

Tract,  cercbro-spinalii  lateralis 


FIG.  152. — The  chief  tracts  descending  to  the  spinal  cord. 

or  Golfs  column  and  the  lateral  fasciculus  cuneatus  or  Burdock' s  column — a  demarcation 
not  emphasized  in  the  lower  part  of  the  cord.  Coil's  column  consists  essentially  of  fibres 
that  come  from  the  lower  segments  of  the  cord  and  is  nothing  more  than  the  continuation  of 
the  laterally  situated  fibres  of  the  lower  segments,  which  during  their  ascent  have  been  pushed 
toward  the  mid-line  by  the  new  increments  of  fibres  entering  at  higher  levels.  Or,  we  may 


PATHS  OF  THE  SPINAL  CORD. 


165 


say,  in  the  cervical  region  of  the  cord,  Coil's  column  is  composed  of  fibres  which  ascend 
from  the  lower  parts  of  the  cord,  and  conveys  sensory  fibres  from  the  lower  extremities 
and  the  lower  half  of  the  trunk,  while  Burdach's  column  carries  sensory  fibres  that  enter 
the  spinal  cord  from  the  upper  half  of  the  trunk  and  the  upper  extremities  (Fig.  151). 


Tractns  spino-cerebellares  dor- 
sails  et  ventralis 


Tract,  spin*  thnlamiats  _  .         __   \_  _ 


FIG.   153. — The  chief  tracts  ascending  from  the  spinal  cord. 

The  terminal  arborization  of  the  ascending  fibres  and  of  the  collaterals  occurs  in 
almost  all  parts  of  the  gray  substance  of  the  same  side  of  the  cord  ;  a  small  part  of  the 
fibres  passes,  by  way  of  the  posterior  commissure,  to  the  opposite  side  to  end  within  the 
posterior  horn.  The  short  fibres  and  the  collaterals  of  the  lateral  bundle,  in  particular, 
end  within  the  homolateral  posterior  horn  and  also  within  the  central  field  ;  the  main  fibies 


166  THE   FIBRE-TRACTS. 

and  collaterals  of  the  median  bundle,  which  end  within  the  cord,  arborize  around  the  cells 
of  Clarke's  column,  the  cells  of  the  intermediate  zone  and  the  anterior  horn  cells.  The 
collaterals  from  the  posterior  column,  which  break  up  around  the  anterior  horn  cells, 
constitute  the  reflex  collaterals. 

The  descending-  branches  of  the  fibres  of  the  posterior  column,  entering  the  root- 
zone  medial  to  the  posterior  horn,  caudalward  form  a  bundle  that  in  cross-section  appears 
comma-shaped.  The  fibres  of  this  field,  the  comma  bundle  of  Schultze,  after  a  short 
course  enter  the  gray  substance. 

In  addition  to  the  chief  fibres,  within  the  posterior  column  are  others  which  arise 
in  the  posterior  horns  of  the  cord,  as  the  axones  of  association-cells.  They  course  within 
the  ventral  part  of  the  posterior  column  and  in  cross-section  appear  as  the  ventral  field 
(Fig.  149). 

Finally  to  be  noted  are  fibres  that  extend  from  the  cervical  region  as  far  as  the 
conus  terminalis.  In  the  upper  regions,  they  lie  dorsally  at  the  periphery  of  the  poste- 
rior column,  more  within  the  area  of  Coil's  column  ;  farther  below,  they  migrate  toward 
the  septum  posterius  and,  within  the  sacral  region,  in  cross-section  appear  as  a  small 
medial  oval  field.  This  part  has  been  termed  the  oval  bundle  of  the  posterior  column. 
It  corresponds  to  the  bandelette  me'diale  of  Gombault  and  Philippe,  the  dorso-medial 
sacral  field  of  Obersteiner  and  the  tractus  cervico-lumbalis  dorsalis  of  Edinger. 

The  chief  tracts  descending  to  and  ascending  from  the  spinal  cord  are  represented  in 
Figs.  152  and  153. 

THE  MEDULLA  OBLONGATA. 

The  medulla  oblongata  forms  the  transition  from  the  spinal  cord  to  the  brain.  The 
intimate  make-up,  relatively  simple  within  the  spinal  cord,  at  the  same  time  undergoes 
manifold  modifications.  Not  only  does  the  gray  substance  change  its  form,  but  new 
masses,  large  and  small  nuclei,  appear.  Coincidentally,  a  rearrangement  of  certain  systems 
of  the  white  substance  occurs,  certain  tracts  of  fibres  are  suppressed  and  new  ones  make 
their  appearance.  Almost  each  cross-section  presents  a  different  picture.  It  would  carry 
us  too  far  to  follow  accurately  at  this  place  the  structure  of  the  medulla  oblongata  in 
its  topographical  relations,  as  shown  in  transverse  sections.  The  study  of  the  fibre- 
tracts  in  the  brain  and  spinal  cord,  without  the  use  of  serial  sections,  is  impossible ; 
and  especially  the  study  of  the  fibre-tracts  in  the  medulla  oblongata  offers  difficulties 
as  nowhere  else. 

The  reader  is  particularly  referred,  therefore,  to  Part  III  of  this  book,  in  which,  by 
means  of  the  drawings  of  serial  sections,  the  most  important  tracts  may  be  followed 
through  the  entire  brain-stem.  By  means  of  these  microscopical  pictures  and  the  assist- 
ance of  the  following  diagrammatic  figures,  the  reader  will  be  able  to  find  his  way. 

In  the  consideration  of  the  morphology,  the  most  important  gray  masses  of  the 
medulla  oblongata  have  been  mentioned  ;  we  may  limit  ourselves,  therefore,  to  the  pres- 
entation of  the  connection  of  these  gray  masses  with  other  parts  of  the  central  nervous 
system.  Then,  as  supplementary,  the  origin  of  the  cerebral  nerves  will  be  more  closely 
considered,  since,  as  pointed  out  in  the  morphological  section,  the  nuclei  of  most  of  the 
cerebral  nerves  are  situated  along  the  floor  of  the  fourth  ventricle. 


PATHS   OF  THE   MEDULLA    OBLONGATA. 


167 


We  proceed  most  advantageously,  if  we  trace  upward  the  fibre-systems  of  the  white 
substance  of  the  cord,  which  were  described  in  the  preceding  section.  At  the  same  time, 
we  will  ignore  the  fibre-systems,  which  come  from  the  brain  and  descend  in  the  cord  and 
have  been  mentioned  repeatedly  in  previous  sections. 

Let  us  first  follow  the  tract  of  the  posterior  column.  The  fibres  of  Burdach's  and 
of  Coil's  column  end  in  the  nuclei  of  the  posterior  column,  that  is,  within  the  nucleus 
fasciculi  cunati  and  the  nucleus  fasciculi  gracilis.  From  these  nuclei  further  tracts  are 
developed,  of  which  one  in  particular,  the  tract  establishing  relations  between  the  pos- 


Lemniscus  medial ts 


Tractus  spino-ihalamicus 


Nucl.  funic.  post. 


FlG.  154. — Posterior  column  tract,  medial  fillet  and  tegmental  tract. 


terior  column  nuclei  and  the  thalamus,  now  concerns  us.  The  fibres  pass,  as  the 
axones  of  the  cells  within  the  cuneate  and  gracile  nuclei,  in  ventrally  directed  courses, 
the  fibrae  arcuatae  internae,  toward  the  mid-line,  where  by  decussation  they  form  the 
raphe.  After  crossing,  the  fibres  assemble  close  to  the  mid-line  and,  turning  upward,  pass 
longitudinally  to  higher  levels.  The  field  so  formed  is  known  as  the  interolivary  stratum, 
on  account  of  its  position  between  the  two  inferior  olivary  nuclei.  The  fibre-bundles 
can  be  traced  through  the  pons  and  the  mid-brain  as  far  as  the  thalamus,  where  they 
end  in  the  nucleus  lateralis  and  in  the  centrum  medianum.  This  is  the  path  usually 
called  the  medial  fillet  or  lemniscus  medialis ;  it  is  also  known  as  the  tractus  bulbo- 
thalamicus. 


1 68 


THE    FIBRE-TRACTS. 


Tractvs  spino-thalamicus . 


FIG.  155. — Course  of  the  medial  fillet. 


THE   MEDIAL   FILLET. 


169 


The  medial  fillet  is  not  composed  exclusively  of  fibres  which  come  from  the  nuclei 
of  the  posterior  columns.  During  its  course  through  the  medulla  oblongata,  the  fillet-tract 
is  augmented  by  the  fibres  from  the  spinal  cord,  which  we  have  studied  as  the  tractus 


N.ruber 


Tractus  cerebello-tegmentalis 


Fibrae  arcuatae  externae  donates 


Nitcl.  gracilis  et  cuneatus 


fibrae  arctiatae  internae, 
ontinued  as  fibrae  arc 
vent?  ales  externae 


Tract,  spino-cerebellaris 
dorsalis  (Flechsig) 


FIG.   156. — Formation  of  the  restiform  body. 

spino-thalamicus.  In  addition,  as  will  be  pointed  out  later,  fibres  come  from  the  terminal 
nuclei  of  the  cranial  nerves.  All  these  contributions  collectively  constitute  the  medial 
fillet,  which  ends  within  the  thalamus. 

Still  other  tracts  arise  from  the    posterior  column  nuclei    and    unite  the  latter  with 
the  cerebellum.      Some    of   these  fibres   pass,  as    do   the  above-mentioned  fibrae  arcuatae 


170  THE   FIBRE-TRACTS. 

internae,  first  towards  the  mid-line  and  there  cross.  They  course,  however,  not  longitu- 
dinally within  the  interolivary  stratum,  but  pass  ventrally  along  the  raphe  as  far  as  the 
anterior  medial  fissure,  then  around  the  pyramids  and  the  olives  as  the  fibrae  arcuatae 
externae  ventrales,  and  continue  as  constituents  of  the  restiform  bodies  to  the  cerebellum. 
Other  fibres  issue  dorsally  from  the  posterior  column  nuclei  and  pass  directly  to  the 
corpus  restiforme  as  the  fibrae  arcuatae  externae  dorsales  (Fig.  156).  Perhaps  these  are 
joined  by  direct  fibres  from  the  posterior  column  tracts. 

The  constitution"  of  the  corpus  restiforme,  or  the  inferior  cerebellar  peduncle,  may 
now  be  considered.  Although  the  tracts  ascending  to  the  cerebellum  from  the  spinal 
cord  and  the  medulla  are  the  principal  factors  in  its  make-up,  additional  paths  of  especial 
importance  are  contributed  to  the  restiform  body  by  the  fibre-bundles  from  the  vestibular 
nerve  and  its  end-nucleus. 

The    restiform    body    consists  of   two  chief  parts,  a  lateral  and  a  medial  division. 

The  lateral  division  is  formed  by  the  following  fibre-bundles: 

a.  The   tracius   spino-cerebellaris    dorsalis   or   direct   cerebellar  tract.     Although  the 
tractus  spino-cerebellaris  ventralis  or  Gowers'   bundle  likewise  passes  to  Che  cerebellum,  it 
reaches  the  latter,  not  by  way  of  the   restiform   body,  but,  farther  above,  in  conjunction 
with  the  superior  cerebellar  peduncle  (Figs.   157  and  178). 

b.  The  fibres  from  the  nucleus  gracilis  and  cuneatus  of  the  same  and  of  the  opposite 
side:  fibrae   arcuatae   externae  dorsales  et  ventrales,  as    well    as    direct    fibres    from    the 
posterior  column. 

c.  A  few  fibres  from  the  nuclei  arcuati  or  pyramidal  nuclei. 

d.  Fibres  from  the  lateral  column  nuclei. 

e.  Fibres  from  the  inferior  olivary  nucleus. 

The  last  fibres — tractus  olivo-cerebellaris — contribute  the  chief  bulk  of  the  lateral 
division  of  the  peduncle.  They  arise  in  largest  part  from  the  contralateral  olive,  a  few 

fibres  coming  also  from  the  olive  of  the 
same  side,  and  end,  as  do  the  other  fibres 
of  the  lateral  division,  within  the  cortex 
of  the  worm. 

The  medial  division  consists  of  two 
chief  varieties  of  fibres: 

a.  One  set  of  fibres  is  the  sensory 
root-fibres  of  certain  cerebral  nerves,  as  the 
trigeminus  and  the  vestibular,  which  pass 

FIG.    157. —  Schematic   representation    of    the    course    of         ,.  ,  u    11  j  i 

the  tractus  spino-cerebellaris  dorsalis  and  ventralis.  dlrect    to     the     Cerebellum     and    Constitute   the 

direct  sensory  cerebellar  tract  of  Edinger. 

b.  The  other  fibres  connect  the  nuclei  of  the  sensory  cerebral  nerves  with  the  cere- 
bellum; among  these  connections,  those  of  the  vestibular  nucleus  with  the  cerebellum 
deserve  special  notice. 

The  termination  of  the  fibres  of  both  sets  is,  in  largest  part,  within  the  nucleus 
tegmenti  of  the  cerebellum.  Fibres  pass  also  in  the  opposite  direction  from  the  nucleus 
tegmenti  or  roof  nucleus  to  the  end-nuclei  of  the  sensory  nerves,  Deiters'  and  Bechterew's 


THE   REST1FORM    BODY.  171 

nuclei  especially  receiving  such  fibres.  These  bundles,  that  bring  the  nuclei  of  the  sensory 
cerebral  nerves  into  relation  with  the  cerebellum,  constitute  the  tractus  nucleo-cerebellaris. 
The  latter  forms  an  indirect  sensory  cerebellar  tracf,  in  contrast  to  the  above-mentioned 
direct  one.  Fibres  from  the  cerebellum  also  pass  caudally  to  the  medulla  oblongata  by 
way  of  the  restiform  body. 

The  tractus  cerebello-bulbaris  or  fastigio-bulbaris  is  a  descending  bundle,  which 
proceeds  especially  from  the  nucleus  tecti  of  the  opposite  side  and,  perhaps,  also  from 
the  nucleus  dentatus.*  The  bundle  is  known  also  as  the  tractus  uncinatus  (Russel-Thomas). 
The  fibres  pass  above  the  superior  cerebellar  peduncle  and,  in  their  farther  course,  reach 
the  medial  division  of  the  restiform  body.  Their  termination  is  partly  within  Deiters', 
nucleus  and  partly,  farther  caudalward,  within  certain  nuclei  of  the  medulla  oblongata, 
along  with  collaterals  to  the  motor  nuclei  of  cerebral  nerves — V,  VII  and  X.  Atten- 
tion has  been  called  to  these  paths,  as  the  tractus  cerebello-tegmentales  bulbi,  when 
considering  the  chief  connections  of  the  cerebellum.  Such  cerebellofugal  tracts  proceed 
from  the  restiform  body,  with  the  fibrae  arcuatae  externae  ventrales,  to  the  olive  and 
the  pyramid  and  ascend  within  the  raphe  to  the  formatio  reticularis  of  the  medulla 
oblongata. 

The  inferior  olivary  nucleus,  as  we  have  seen,  gives  off  a  robust  fibre-bundle,  which 
passes  to  the  corpus  restiforme  of  the  opposite  side  and,  thence,  to  the  cerebellum.  A 
small  number  of  fibres,  on  the  other  hand,  arise  within  the  cerebellar  cortex  and  descend 
to  the  opposite  olive.  The  olivary  nucleus  possesses  still  further  connections.  Thus, 
from  the  ascending  tractus  spino-cerebellares  collaterals  are  sent  to  the  inferior  olivary 
nucleus,  while  other  fibres  are  received  from  the  tractus  spino-olivaris  or  Helweg's  tri- 
angular tract,  as  well  as  from  the  tractus  thalamo-olivaris.  By  means  of  the  last-mentioned 
path,  impulses  from  the  thalamus,  and  also  from  the  cerebral  cortex  by  way  of  the 
thalamus,  are  carried  to  the  inferior,  olive  and,  by  way  of  the  olivo-cerebellar  tract,  to  the 
cerebellum  (Fig.  145). 

The  medial  fillet  or  interolivary  stratum  appears  in  cross-sections  of  the  medulla 
as  a  field,  that  lies  between  the  two  olives  at  the  sides  of  the  raphe  (see  Part  III). 
Dorsally,  as  the  apex  of  this  field,  the  posterior  longitudinal  bundle  or  the  fasciculus 
longitudinalis  medialis  is  seen  as  a  small  bundle  of  longitudinally  coursing  fibres.  It  will 
be  considered  more  fully  in  connection  with  the  vestibular  nerve  (page  182).  Lateral  to 
the  interolivary  stratum  and  dorsal  to  the  olive,  a  field  spreads  out  which,  in  addition 
to  numerous  scattered  nerve-cells,  contains  longitudinally  coursing  nerve-fibres.  This 
area,  the  upward  continuation  of  the  formatio  reticularis  of  the  spinal  cord,  is  known 
as  the  association  field  of  the  medulla  oblongata.  The  formatio  reticularis  extends  far 
up  into  the  mid-brain  and  contains  numerous  connecting  paths,  of  longer  or  shorter 
course,  by  which  manifold  relations  are  established  between  certain  nerve-nuclei.  It  is 
probable  that  within  this  formatio  reticularis  run  those  association  fibres  which  unite  the 
nuclei  of  the  vagus,  facial  and  phrenic  nerves  for  coordinated  activity  during  respiration. 
Attention  must  be  called  again  to  the  tractus  spino-thalamicus,  which,  composed  of  axones 
of  posterior  horn  cells,  crosses  to  the  opposite  lateral  column  of  the  spinal  cord  and,  without 
further  interruption,  passes  to  the  thalamus.  This  tract  is  an  important  sensory  pathway, 
supplementary  to  the  median  fillet. 


172 


THE   FIBRE-TRACTS. 


THE  CEREBRAL  NERVES. 

NERVUS    OLFACTORIUS. 

The  first  member  of  the  conventional  series  of  twelve  cranial  nerves,  the  nervus 
olfactorius,  is  represented  by  the  short  peripheral  paths,  the  fila  olfactoria,  connecting  the 
olfactory  mucous  membrane  with  the  glomeruli  within  the  bulbus  olfactorius.  Since  the 
structures  formerly  described  as  the  first  cerebral  nerve,  the  olfactory  bulb  and  tract,  are 
parts  of  the  rhinencephalon,  their  consideration  falls  properly  with  that  of  the  brain. 
They  have  been  discussed  under  the  Fibre- Paths  of  the  Rhinencephalon  (page  144),  to 
which  the  reader,  therefore,  is  referred. 


Left 


Right 


NERVUS   OPTICUS. 

The  fibres  of  the  optic  nerve  arise  within  the  retina  and  are  the  axones  of 
the  ganglion  cells  located  within  the  ganglion-cell  layer  of  the  nervous  tunic  of  the  eye. 
They  extend  to  the  chiasm.  Here,  one  part  of  the  fibres  passes  to  the  tractus  opticus 

of  the  opposite  side,  the  other  part  passes 
direct  to  the  tract  of  the  same  side.  The 
fibres  end  within  the  corpus  geniculatum 
laterale,  the  pulvinar  and  the  superior 
colliculus ;  these  end-stations  constitute 
primary  visual  centres.  From  the  lateral 
geniculate  body  and  the  pulvinar,  fibres 
pass  through  the  hindmost  part  of  the 
posterior  limb  of  the  capsula  interna  to 
the  secondary  or  cortical  visual  centres 
within  the  cortex  of  the  cuneus,  thereby 
forming  the  optic  radiation  of  Gratiolet. 
Fibres  also  pass  in  the  opposite  direction 
from  the  cortical  visual  centre  to  the 
primary  centres.  It  must  be  noted  fur- 
ther, that  fibres  exist,  which  arise  within 
the  primary  centres  and  end  within  the 
retina. 

The  visual  fibres  proper  terminate 
within  the  corpus  geniculatum  laterale 
and  the  pulvinar  thalami  and  probably  do 

not  invade  the  superior  colliculus  of  the  corpus  quadrigeminum,  at  least  in  the  higher 
vertebrates.  The  optic  fibres  which  pass  to  the  superior  colliculus  are  concerned  with  a 
special  duty.  Stimuli  carried  by  these  fibres  to  the  superior  colliculus  are  transferred  to  the 
deeper  lying  oculomotor  nucleus,  resulting  in  the  liberation  of  the  pupillary  reflex.  These 
optic  fibres  ending  in  the  superior  colliculus  are  known,  therefore,  as  pupillary  fibres. 


Optic 
radiatii 


FIG.  158. — The  path  of  visual  impulses. 


CEREBRAL   NERVES.  173 

The  pupillary  reflex  consists  in  a  contraction  of  the  pupil  in  response  to  the 
entrance  of  light.  The  reaction  is  exhibited  by  both  eyes  ;  that  is,  when  the  light 
falls  on  only  one  eye,  the  contraction  occurs  not  only  in  the  stimulated  eye  (direct 
reaction),  but  also  in  the  other  eye  (consensual  reaction). 


FIG.  159.— The  course  of  the  visual  path. 

The  intercentral  paths  of  the  pupillary  reflex  are  not  positively  established,  although 
It  may  be  regarded  as  certain,  that  the  entire  reflex-tract  is  made  up  of  a  sequence 
including  several  neurones.  It  may  be  assumed  that  the  stimulus  passes : 

a.  From  the  retina  to  the  superior  colliculus  ; 

b.  From  the  superior  colliculus  to  the  nucleus  of  the  oculomotor  nerve ; 

c.  From  the  oculomotor  nucleus  to  the  ciliary  ganglion  ; 

d.  From  the  ganglion  ciliare  to  the  sphincter  pupillae  muscle. 

Since  the  illumination  of  one  eye  causes  uniform-  contraction  of  both  pupils,  the  re- 
flex being,  therefore,  homo-  and  heterolateral,  it  follows  that  the  impulse  from  one  colliculus 
must  be  transferred  to  both  oculomotor  nuclei,  or  one  nucleus  must  be  able  to  stimulate 


>74 


THE   FIBRE-TRACTS. 


Ganglion 
citiare 


both  the  right  and  left  sphincter  pupillae  muscles.  The  particular  fibre-bundle,  by  means  of 
which  the  impulse  is  transferred  from  the  superior  colliculus  to  the  oculomotor  nucleus,  is  not 
definitely  determined.  In  Fig.  160,  the  reflex  path  is  schematically  represented,  with  the  as- 
sumption, that  the  fibres  proceeding  from  the  superior  colliculus  reach  both  oculomotor  nuclei. 
A  knowledge  of  the  course  of  the  fibres  of  the  optic  nerve,  particularly  their  semi- 
decussation,  supplies  the  explanation  of  one  of  the  most  important  disturbances  of  vision — 
hemiopia  (half  seen)  or  hemiariopsia  (half  not  seen).  If  the  conduction  of  one  optic  tract, 
for  example  the  left,  be  interrupted  by  a  lesion,  the  stimuli  coming  from  the  left  retinal 
halves  of  the  two  eyes  can  no  longer  be  transmitted  to  the  cortical  centres  in  the  left 
hemisphere,  the  right  halves  of  the  visual  fields  are  lost  and  only  the  left  halves  of  fixed 
objects  are  still  seen  (Fig.  158).  This  condition  is  spoken  of  as  homolateral  or  homony- 
mous  hemianopsia  or  hemiopia.  Lesion  of  the  left  tractus  leads  to  right-sided  homonymous 
hemianopsia  or  to  left-sided  hemiopia  ;  lesion  of  the  right  tract  leads  to  left-sided  homonymous 

hemianopsia  or  to  right-sided  hemiopia. 
Homonymous  hemianopsia  follows,  of 
course,  not  only  lesion  of  the  tractus 
opticus,  but  also  lesion  of  the  secondary 
paths  connecting  the  primary  and  sec- 
ondary centres,  that  is  within  the  optic 
radiation,  or  lesion  of  the  cortical  centre. 
In  relation  to  the  diagnosis  of  the  seat 
of  the  lesion,  the  pupillary  reaction  pos- 
sesses a  certain  significance.  If  in  ho- 
monymous hemianopsia  the  light-reflex 
is  lost  when  the  insensitive  half  of  the 
retina  is  illuminated,  the  seat  of  the  lesion 
is  the  tractus  (Wernicke's  hemianopsic 
pupillary  rigidity  or  hemiopic  pupillary 
reaction).  If,  on  the  contrary,  the 
light-reflex  of  the  pupil  is  intact,  then 
the  lesion  lies  hjgher,  for  example,  in 

the  internal  capsule  or  in  the  cerebral  cortex.  In  the  majority  of  cases  of  homonymous  hemian- 
opsia, we  have  to  do  with  tumors  of  the  occipital  lobe,  more  rarely  with  lesions  of  the  tractus 
opticus.  Complications  associated  with  hemianopsia,  such  as  hemiplegia,  hemiparesis,  con- 
tractions and  aphasic  disturbances  (with  right-sided  hemianopsia),  must  also  be  borne  in  mind. 
The  same-sided  or  homonymous  hemianopsia  is  the  opposite  of  the  heteronymous 
hemianopsia,  which  occurs  more  rarely  than  the  homonymous.  When  the  temporal  halves 
of  both  visual  fields  are  wanting,  such  heteronymous  hemianopsia  is  known  as  temporal 
hemianopsia.  In  such  cases,  the  lesion  is  situated  within  the  chiasm,  either  in  the  middle 
or  in  the  anterior  or  the  posterior  angle  of  the  chiasm,  whereby  the  decussating  fibres  are 
involved.  Temporal  hemianopsia  is  observed,  for  example,  in  acromegaly,  in  which  the 
enlargement  of  the  hypophysis  cerebri  concurrently  affects  the  chiasm.  When  the  nasal 
halves  of  both  visual  fields  are  wanting,  the  condition  is  spoken  of  as  nasal  hemianopsia 
and  is  produced  by  involvement  of  the  uncrossed  fibres,  as  when  the  chiasm  is  subject  to 
pressure  on  both  sides  in  the  lateral  angle  by  enlarged  carotids. 


Corte. 
(visual  centre^) 


N.  oculomotorius 

FIG.   160. — The  pupillary  reflex  path. 


Cortex 
>tal  centre) 


CEREBRAL   NERVES. 


175 


NERVUS   OCULOMOTORIUS. 

The  oculomotor  nerve  arises  in  the  nucleus  nervi  oculomotorii,  which  lies  in  the  region 
of  the  superior  colliculus,  ventral  to  the  aquaeductus  Sylvii,  within  the  floor  of  the  central  gray 
substance  (Figs.  88  and  89).  The  nucleus  consists  of  a  medially  placed  medial  nucleus  and 


Accommodation 


FIG.  161. — The  deep  origins  of  the  motor  cerebral  nerves. 

a  pair  of  large-celled  lateral  nuclei.  The  nerve  conveys  fibres  which  originate  within  the  me- 
dial nucleus  and  the  lateral  nucleus  of  the  same  and,  in  part,  of  the  opposite  side.  The  fibres 
pass  ventrally  in  laterally  convex  curves  and  emerge  from  the  brain-stem  along  the  sulcus  nervi 
oculomotorii  on  the  medial  surface  of  the  pedunculus  cerebri.  The  voluntary  innervation  of 
the  nucleus  proceeds,  as  in  the  case  of  all  the  motor  cere- 
bral nerves,  from  the  cerebral  cortex.  The  entire  path  of 
conduction  includes: 

Hevatorpalpebrae 

a.  The  central  neurone — cerebral  cortex  to  nucleus;         N^~^^  A  fP/^^Re<tus  superior 

7       T^I  •*  i  t  1  -it  vv\N  ^^   l/j\/  RectuS  i 

b.  The    peripheral    neurone  —  nucleus,     peripheral  XVv|||  pv/ Ob,^uus  inferior 

nerve,    muscle.  Pi  W  fJ  Rectus  inferior 

It  must  be  pointed  out,  however,  that  the  course  of 
the  central  path  is  not  yet  known;  probably  the  path  is 
composed  of  several  neurones.  Likewise,  uncertainty  exists 
regarding  the  location  of  the  cortical  centres,  which  have  been 
variously  assumed  as  lying  within  the  gyrus  angularis,  the 

occipital  lobe  or  the  frontal  lobe.  The  centre  for  voluntary  eye-movements,  however,  is  quite 
generally  regarded  as  including  the  posterior  part  of  the  second  or  middle  frontal  convolution. 
Investigations  have  shown,  that  the  entire  oculomotor  nucleus  is  made  up  of  certain  groups  of 
cells,  of  which  a  particular  group  always  gives  origin  to  the  fibres  for  a  particular  muscle.  Con- 
cerning these  special  subdivisions  of  the  nucleus,  however,  we  shall  not  enter  more  fully,  for 
the  reason  that  these  relations,  as  yet,  have  been  by  no  means  definitely  established.  Fig.  162 


N-IIC.  n.  trochlenris 


FIG.  162. — Subsidiary  nuclei  of  the  ocu- 
lomotor nucleus.     (Bernheimer.) 


I76 


THE   FIBRE-TRACTS. 


represents  the  individual  cell-groups,  according  to  the  investigations  of  Bernheimer  on 
monkeys.  In  the  middle  is  the  medial  chief  nucleus,  on  either  side  the  lateral  chief  nucleus 
with  its  various  subdivisions,  and,  medial  from  the  latter,  the  small  lateral  nucleus,  which 
is  also  known  as  the  nucleus  of  Edinger-  Westphal. 

NERVUS   TROCHLEARIS. 

The  trochlear  nerve  has  its  origin  in  the  nucleus  nervi  trochlearis,  which  is  located 
in  the  caudal  prolongation  of  the  oculomotor  nucleus  in  the  region  of  the  inferior  colliculus. 
The  fibres  of  the  fourth  nerve  pass  dorsally,  cross  in  the  velum  medullare  anterius  and 
emerge  from. the  brain-stem  behind  the  quadrigeminal  bodies,  on  either  side  of  the  frenu- 
lum  veli  medullaris  anterioris.  As  in  the  case  of  the  oculomotor  nervi,  the  path  includes: 

a.  The  central  neurone — from  cerebral  cortex  to  nucleus; 

b.  The  peripheral  neurone — nucleus,   peripheral  nerve,   muscle. 

NERVUS   ABDUCENS. 

The  nucleus  of  the  abducens  nerve  lies  in  the  floor  of  the  fourth  ventricle  and  in 
the  colliculus  facialis.  The  emergent  fibres  of  the  sixth  nerve  pass  ventrally  and  leave 
the  brain-stem  at  the  posterior  border  of  the  pons.  The  path  includes: 

a.  The  central  neiirone — from  cerebral  cortex  to  nucleus; 

b.  The  peripheral  neurone — nucleus,   peripheral  nerve,   muscle. 

NERVUS   TRIGEMINUS. 

Here  a  motor  part  and  a  sensory  part  are  to  be  distinguished  (Figs.  163  and  164). 

I.    Motor  Portion.     The  central  neurone  takes  origin  in  the  cerebral  cortex  of  the 

lower   third  of   the  central    convolutions,  passes  with  the  pyramidal   tracts  downward  and 


Nucleus  radicis  dcscendentis  N.  trigemhtt 


2KX 


FIG.  163.  —  Course  of  the  trigeminus,  vagus  and  glossopharyngeus  within  the  brain-stem. 


CEREBRAL   NERVES. 


177 


ends  in  the  chief  motor  nucleus,  within  the  dorso-lateral  part  of  the  tegmentum  of  the 
pons.  The  peripheral  neurone  arises  within  this  motor  nucleus,  the  motor  root  of  the 
nerve  receiving  also  fibres  from  the  nucleus  of  the  opposite  side.  The  fibres  emerge  from 
the  pons  as  the  portio  minor  nervi  trigemini  and  pass  to  the  muscles.  A  small  part  of 
the  motor  root  arises  from  small  cells,  which  lie  lateral  to  the  Sylvian  aqueduct  within 
the  region  of  the  corpora  quadrigemina  and  constitute  the  nucleus  radicis  descendentis 


Xam.I 


Central  trigeminal  trac 


•y  terminal  nucleus 


Descending  sensory  trigeminal 

roof  an(i  fa  ending  in  sensory 

nucleus 


FIG.   164. — Course  and  relations  of  the  root-fibres  of  the  trigeminal  nerve. 

nervi  trigemini.  This  group  of  cells  joins  caudally  the  cell-area  of  the  locus  caerulus. 
The  fibres  arising  from  these  cells,  after  giving  off  collaterals  to  the  chief  motor  nucleus, 
pass  outward  with  the  other  peripherally  directed  processes  of  the  motor  neurones. 

2.  Sensory  Portion.  The  origin  of  the  sensory  part  of  the  trifacial  nerve  lies 
within  the  ganglion  Gasseri.  The  axones  of  the  unipolar  ganglion  cells  of  this  ganglion 
divide  into  two  branches.  One  of  these  extends  peripherally  as  the  peripheral  nerve, 
the  other  passes  centrally,  enters  the  pons  as  the  portio  major  nervi  trigemini,  and  runs 
to  the  sensory  end-nucleus  of  the  trigeminus,  close  to  the  motor  nucleus.  Here  each 


1 78  THE   FIBRE-TRACTS. 

fibre  divides  into  an  ascending  and  a  descending  branch.  The  ascending  branch  ends 
within  the  sensory  nucleus  within  the  pontile  tegmentum.  The  descending  branch  ends, 
after  giving  off  numerous  collaterals,  in  a  nucleus  that  is  nothing  more  than  the  caudal 
prolongation  of  the  sensory  nucleus.  The  descending  branches  form  collectively  the 
tractus  spinalis  nervi  trigemini;  the  gray  substance  in  which  this  path  ends,  constitutes 
the  nucleus  tractus  nervi  trigemini.  The  descending  tract,  as  well  as  the  nucleus,  can 
be  followed  downward  as  far  as  the  cervical  cord,  the  nucleus  being  identical  with  the 
substantia  gelatinosa  Rolandi  capping  the  posterior  horn.  From  the  sensory  end-nucleus 
arises  the  //  neurone.  The  fibres  pass  towards  the  mid-line,  giving  off  collaterals  to  the 
nucleus  of  the  facial  nerve,  cross  to  the  fillet-tract  of  the  opposite  side,  there  turn  upward 
and  run  forward  (partly  within  the  medial  fillet  and  partly  as  a  more  laterally  placed 
special  ascending  bundle),  and  later  enter  the  thalamus  with  the  medial  fillet.  Finally, 
a  ///  neurone  succeeds  the  second  one,  thus  linking  the  thalamus  with  the  cortical  sensory 
area.  Still  to  be  mentioned  are  sensory  fibres,  which  pass  direct  to  the  cerebellum  as 
constituents  of  the  direct  sensory  cerebellar  tract;  further,  fibres  which  pass  from  the 
sensory  end-nucleus  to  the  cerebellum  as  constituents  of  the  tractus  nucleo-cerebellaris. 

NERVUS   FACIALIS  AND   NERVUS   INTERMEDIUS   WRISBERGI. 

The  facial  nerve  arises  in  the  nucleus  nervi  facialis,  which  lies  within  the  ventral 
area  of  the  pontile  tegmentum,  ventro-lateral  to  the  abducent  nucleus.  The  fibres  spring- 
ing from  the  nucleus  first,  proceed  dorsally,  pass  around  the  nucleus  of  the  abducent 
nerve — the  facial  knee  and  the  colliculus  facialis — then  course  ventrally  and  emerge 
from  the  brain-stem  at  the  posterior  border  of  the  pons,  lateral  to  the  olive.  The  volun- 
tary innervation  of  the  nucleus  is  effected  by  fibres,  which  start  from  the  lower  third  of 
the  precentral  convolution,  pass  through  the  internal  capsule  (knee),  then  through  the 
cerebral  peduncle  to  the  pons,  and,  finally,  to  the  homo-  and  the  contralateral  facial 
nucleus.  The  path  includes,  as  in  the  case  of  the  other  motor  nerves: 

a.  The  central  neurone — from  cerebral  cortex  to  nucleus; 

b.  The  peripheral  neurone — nucleus,   peripheral  nerve,   muscle. 

The  facial  nucleus  resembles  the  oculomotor  nucleus  in  including  a  number  of 
different  groups  of  cells,  in  which,  in  the  first  place,  we  distinguish  two  chief  groups,  an 
upper  and  a  lower  facial  nucleus.  The  upper  contains  cells,  whose  axones  form  collec- 
tively the  superior  facial  branch;  the  lower,  cells  whose  axones  form  the  inferior  facial 
branch.  Moreover,  the  superior  facial  nucleus  receives  its  innervation  from  the  motor 
centres  of  both  hemispheres,  a  fact  of  clinical  significance.  This  bilateral  innervation 
explains  why,  in  central  facial  paralysis,  the  muscles  supplied  by  the  upper  facial  division 
are  not  involved  in  the  paralysis,  since  innervation  is  still  provided  by  the  unaffected 
central  neurones  of  the  other  hemisphere.  In  peripheral  facial  palsy,  on  the  contrary, 
all  the  muscles  supplied  by  both  the  upper  and  lower  facial  are  paralyzed. 

The  nervus  intermedius  Wrisbergi  (the  nerve  of  Sapolini,  by  whom  it  was 
regarded  as  the  thirteenth  cerebral  nerve)  is  a  mixed  nerve,  which  accompanies  the 
facial  and  continues  as  the  chorda  tympani.  The  motor  fibres  take  origin  in  a  small  cell' 
group  lying  dorso-medial  to  the  facial  nucleus .  The  sensory  fibres  arise  within  the  gan- 
glion geniculi.  The  axones  arising  from  the  cells  of  this  ganglion  divide  into  two  branches. 


CEREBRAL   NERVES. 


179 


One  of  these  passes  peripherally  and  forms,  after  joining  the  motor  fibres,  the  peripheral  nervus 
intermedius,  that  continues  as  the  chorda  tympani.  The  other  branch  passes  centrally, 
enters  the  brain-stem  and  ends  within  the  nucleus  tractus  solitarii  as  part  of  the  gustatory 
path  (page  189). 

NERVUS   ACUSTICUS. 
The  acoustic  nerve  consists  of  two  parts,  the  nervus  cochleae  and  the  nervus  vestibuli. 


Cortex 
(auditory  centre) 


-  Corpus  geniculaf. 

medial? 

Corpus  quadrigemin. 
post. 


i.    NERVUS  COCHLEAE. 

This  nerve  takes  origin  within  the  ganglion  spirale  cochleae.  The  peripherally 
directed  fibres  of  these  bipolar  ganglion  cells  run  to  the  auditory  cells  within  the  organ 
of  Cord;  the  centrally  directed  fibres  enter  the  brain-stem  and  end  in  two  nuclei.  The 
latter  are  the  nucleus  ventralis  nervi  cochleae,  situated  ventral  and  lateral  to  the  corpus 
restiforme,  and  the  nucleus  dorsalis  nervi  cochleae,  or  tubercuhim  acusticum,  which  lies 
dorsally,  although  connected  with 
the  ventral  nucleus.  The  impulses 
carried  by  these  peripheral  neu- 
rones are  conveyed  to  the  higher 
levels  by  the  central  path  including: 

a.  Neurones  passing  from  the 
nucleus  ventralis    to  the  mid-line, 
forming  the  corpus  trapezoides.   The 
path  is  augmented  by  fibres  from  the 
superior  olive  and  from  the  nucleus 
of   the    corpus   trapezoides.     After 
crossingthe  mid-line, some  fibres  end 
within  the  superior  olivary  nucleus, 
while  others  are  joined  by  fibres  from 
the  nucleus  of  the  corpus  trapezoides 
and  from  the  superior  olive  of  the 
side  on  which  the  path  now  runs. 

The    fibres   form  collectively 
the  lateral  Jillet  or  lemniscus  later- 

alis,  which  ends  within  the  corpus  geniculatum  mediale  and,  chiefly  by  collaterals,  within 
the  inferior  colliculus.  Some  fibres  extend  as  far  as  the  superior  colliculus.  The  lateral 
fillet  receives  additional  fibres  from  a  group  of  cells  lying  in  the  midst  of  the  tract, 
known  as  the  nucleus  lemnisci  lateralis. 

b.  Neurones    passing    from    the    nucleus    dorsalis  or  tuberculum  acusticum  over   the 
corpus  restiforme  and,  as  the  superficial  striae  acusticae,  toward  the  mid-line;  thence  cours- 
ing deeply  to  cross  the  raphe  and  reach  the  opposite  superior  olive,  they  join  the  lateral 
fillet  and  finally  end  within  the  corpus  geniculatum  mediale. 

c.  Neurones  arising  within  the  lateral   geniculate   body  and  passing  to  the  auditory 
centre  within  the  cortex  of  the  gyrus  temporalis  superior.      Fibres  also  run  in  the  opposite 
direction,  from    the  auditory    centre    to  the    medial   geniculate   body    and  to  the   inferior 
colliculus. 


Nucleus 

ventralis 


'  Oliva  superior 


Nuc.  corpor.  trapez. 
FIG.  165. — The  auditory  path. 


i8o 


THE    FIBRE-TRACTS. 


N.  cochleae 


FIG.  166. — Course  followed  by  the  auditory  impulses.    * 


CEREBRAL   NERVES. 


181 


2.   NERVUS  VESTIBULI. 

The  vestibular  division  of  the  auditory  nerve  arises  within  the  ganglion  vestibulare 
or  Scarp  a' s  ganglion,  located  at  the  bottom  of  the  internal  auditory  canal.  The  periph- 
erally directed  processes  or  fibres  of  these  ganglion  cells  run  to  the  ampullae,  the 
utricle  and  saccule  of  the  internal  ear;  the  centrally  directed  fibres  enter  the  brain-stem 
and  divide  into  ascending  and  descending  branches.  The  descending  branches  form  a 
descending  vestibular  root  and  -end  within  the  nucleus  nervi  vestibularis  spinalis,  which 
extends  as  far  as  the  posterior  column  nuclei.  The  ascending  branches  end  within  the 
nucleus  medialis,  as  well  as  within  the  lateral  Deiters^  nucleus  and  the  upper  Bechterew*  s 
nucleus.  From  these  end-nuclei,  fibres  pass  to  the  cerebellar  worm  as  constituents  of  the 
tractus  nucleo-cerebellaris.  A  part  of  the  vestibular  fibres  pass  direct  to  the  roof-nucleus 


Direct  sensory 
cerebellar  tract 


Nucleus  Bechterew 


Nucleus  Deiters 
Nucleus  vestibuli   . 


dialis 
FIG.   167. — Path  of  the  impulses  from  the  vestibular  nerve. 

of  the  cerebellum  as  constituents  of  the  direct  sensory  cerebellar  tract,  the  fibres  giving 
off  collaterals  to  Deiters'  nucleus.  The  medial  nucleus  is  brought  into  relation  with  the 
superior  olive  by  means  of  fibres.  Perhaps  fibres  pass  also  to  the  formatio  reticularis  and 
to  the  thalamus. 

In  view  of  its  importance,  the  system  of  Deiters'  nucleus  claims  closer  attention. 
This  nucleus  receives,  on  the  one  hand,  fibres  from  the  roof-nucleus  of  the  cerebellum  ; 
on  the  other  hand,  as  we  have  seen,  Deiters'  nucleus  gives  origin  to  a  fibre-bundle  that, 
as  the  tractus  vestibulo-spinalis,  passes  to  the  spinal  cord. 

Within  the  same  nucleus,  moreover,  also  another  path,  the  posterior  longitudinal 
bundle  or  the  fasciculus  longitudinalis  media/is,  takes  its  origin.  The  fibres  pass  from 
Deiters'  nucleus  toward  the  mid-line,  some  crossing  the  latter  and  then  dividing  into 
ascending  and  descending  branches.  The  ascending  branches  can  be.  followed  upward  as 
far  as  the  oculomotor  nucleus ;  the  descending  branches  pass  to  the  anterior  column  of  the 


182 


THE   FIBRE-TRACTS. 


spinal  cord.  The  posterior  longitudinal  bundle,  however,  does  not  consist  exclusively  of 
fibres  from  Deiters'  nucleus.  Other  fibres  take  origin  in  the  common  nucleus  of  the 
commissura  posterior  and  of  the  fasciculus  longitudinalis  medialis  within  the  forepart  of 


"  :  -•  Tract,  vestib-ulo-spinalis 


FIG.  168. — Origin  and  course  of  the  posterior  longitudinal  bundle. 

the  mid-brain,  in  front  of  the  oculomotor  nucleus.  The  posterior  longitudinal  bundle  may 
be  traced  from  its  nucleus  through  the  mid-brain,  the  pons  and  the  medulla  oblongata 
into  the  spinal  cord,  during  its  course  giving  off  numerous  collaterals  to  the  nuclei  of  the 


CEREBRAL   NERVES. 


183 


nerves  supplying  the  ocular  muscles.  This  bundle  is  of  great  importance.  It  establishes 
relations  of  the  nuclei  of  the  eye-muscles  to  one  another,  among  which  that  of  the 
abducens  to  the  oculomotorius  nucleus  deserves  particular  attention.  Of  especial  impor- 
tance is  the  connection  of  the  abducent  nucleus  with  those  cells  of  the  oculomotor  nucleus, 
from  which  pass  the  fibres  for  the  rectus  internus,  since  the  synergic  function  of  the 


Oliva  superior 


FIG.  169. — Course  of  the  auditory  path.    Connection  of  the  superior  olive  with  the  nucleus  of  the  abducens  (VI)  and,  by  means 
of  the  posterior  longitudinal  bundle,  with  the  nuclei  of  the  other  nerves  (///,  IV)  to  the  ocular  muscles. 


rectus  externus  and  internus,  which  consists  in  the  conjugate  deflection  of  the  eyes  toward 
one  side,  can  be  explained  only  by  the  existence  of  a  direct  or  an  indirect  connection 
between  these  nuclei.  Fig.  168  represents  the  manner  in  which  the  coordinate  action  of 
the  two  muscles  may  be  explained  upon  an  anatomical  basis.  Connection  of  the  abducens 
nucleus  with  that  of  the  oculomotorius,  by  means  of  the  posterior  longitudinal  bundle,  is 


184  THE   FIBRE-TRACTS. 

positively  established.  Further,  that  the  nerve-fibres  for  the  rectus  internus  arise,  in 
greater  part,  from  the  cells  of  the  oculomotor  nucleus  of  the  opposite  side.  On  the  other 
hand,  it  is  still  undecided,  whether  the  efferent  paths  from  the  cortical  centre  for  synergic 
eye-movements  are  first  interrupted  in  a  special  centre  within  the  quadrigeminal  region, 
or  pass  directly  into  the  posterior  longitudinal  bundle.  In  any  event,  this  tract  undergoes 
a  total  or  partial  decussation  before  it  enters  the  posterior  longitudinal  bundle.  In  Fig. 
1 68,  the  path  from  the  cortex  to  the  nucleus  of  the  bundle  is  represented  as  crossed. 
In  tnis  way,  the  explanation  for  the  following  phenomena  is  supplied.  When  a  cortical 
centre  for  eye-movements  is  stimulated,  the  left  one  for  example,  deviation  of  both  eyes 
toward  the  right  occurs.  On  the  other  hand,  in  left-sided  disease  of  the  cerebral  cortex, 
followed  by  paralysis  of  the  right  half  of  the  body,  deflection  of  both  eyes  toward  the 
side  of  the  lesion,  that  is  the  left,  is  frequently  observed,  since,  under  these  conditions, 
the  eye-muscle  nerves  of  the  left  side  functionally  predominate.  "In  lesions  of  the  hemi- 
spheres, if  there  is  conjugate  deviation  of  the  eyes,  the  patient  looks  toward  the  injured 
hemisphere  when  there  is  paralysis,  or  the  limbs  are  contorted  during  a  convulsion" 
(Grasset).  The  diagram  explains,  further,  the  deviation  of  the  eyes,  toward  the  side 
opposite  to  the  seat  of  the  lesion,  frequently  observed  in  diseases  of  the  pons.  For 
example,  if  a  lesion  of  the  posterior  longitudinal  bundle  lies  in  the  vicinity  of  the  right 
abducens  nucleus,  deviation  of  the  eyes  toward  the  right  occurs  in  consequence  of  the 
mastery  by  the  nerves  controlling  the  left  eye-muscles. 

The  posterior  longitudinal  bundle  possesses  further  importance,  since  it  brings  the 
vestibular  apparatus  and  the  cerebellum  into  relation  with  the  nuclei  of  the  eye-muscles 
and  the  spinal  cord,  by  means  of  the  fibres  arising  within  Deiters'  nucleus.  It  unites, 
therefore,  the  centres  concerned  in  maintaining  equilibrium  and  orientation  in  space. 

It  is  to  be  noted,  that,  since  a  connection  between  the  superior  olive  and  the 
abducens  nucleus  exists,  relations  of  the  acoustic  nerve,  that  is  of  the  auditory  path,  with 
the  abducens,  and,  by  means  of  the  posterior  longitudinal  bundle,  with  the  other  nuclei 
of  the  eye-muscle  nerves  may  also  be  established.  These  connections  explain  the  occur- 
rence of  reflex  ocular  movements  in  response  to  auditory  impressions  (Fig.  169). 

NERVUS  GLOSSOPHARYNGEUS  AND  VAGUS. 

i.  Motor  Portion.  The  efferent  fibres  arise  partly  within  the  nucleus  motorius 
dor  satis  nervi  vagi  et  glossopharyngei,  which  lies  in  the  floor  of  the  fourth  ventricle 
lateral  to  the  hypoglossal  nucleus  and  medial  to  the  nucleus  alae  cinereae  ;  the  larger 
part  of  the  fibres,  however,  arises  from  the  cells  of  the  nucleus  ventralis  or  ambiguus, 
which  lies  within  the  formatio  reticularis  dorsal  to  the  dorsal  accessory  olive.  Since  the 
voluntary  innervation  of  the  nucleus  proceeds  from  the  cerebral  cortex,  the  path  includes  : 

a.  The  central  neurone — from  the  cerebral  cortex  to  nucleus; 

b.  The  peripheral  neurone — nucleus,  peripheral  nerve,  muscle. 

The  root-bundles  passing  out  from  the  dorsal  nucleus  are  the  equivalents  of  motor 
preganglionic  sympathetic  fibres  destined  for  the  innervation  of  involuntary  muscle  ;  this 
nucleus,  therefore,  is  also  designated  as  the  sympathetic  motor  nucleus.  The  fibres  pro- 
ceeding from  the  nucleus  ambiguus,  on  the  contrary,  are  for  the  voluntary  muscle  ;  this 
nucleus,  therefore,  is  known  as  the  somatic  motor  nucleus.  The  latter  consists  of  several 


CEREBRAL   NERVES. 


185 


groups  of  nerve-cells,  the  individual  groups  representing  centres  for  the  particular  groups 
of  muscles  innervated  by  the  vagus.  The  positions  of  these  centres  within  the  nucleus, 
however,  are  not  yet  sufficiently  determined. 

2.  Sensory  Portion.  The  efferent  fibres  arise  within  the  ganglion  superius  et 
pctrosum  nervi  glossopharyngei  and  ganglion  jugulare  et  nodosum  nervi  vagi  respectively. 
The  peripherally  directed  branches  form  the  peripheral  sensory  nerves ;  the  centrally 
directed  branches  enter  the  brain-stem  as  the  sensory  root-fibres  and  pass  to  the  end- 
nuclei.  One  part  of  the  fibres  ends  within  the  nucleus  aloe  cinereae,  while  another  part 


Nucleus  tractus 
solitarli 


FIG.  170. — Transverse  section  of  the  medulla  ob- 
longata.  Origin  of  the  IX  and  X  (motor  part)  and  of 
the  XII  nerve. 


Tract,  soli  tar  ins 


PIG.  171. — Origin  of  the  IX  and  X  nerves,  sensory  part. 


forms  a  descending  root,  the  tractus  solitarius,  and  ends  within  the  accompanying  tract  of  gray 
substance,  the  nucleus  tractus  solitarii.  The  central  neurones  arise  within  the  end-nuclei. 
The  fibres  emerging  from  the  end-nuclei  pass  toward  the  mid-line  and  the  interolivary  layer, 
thence  with  the  medial  fillet  to  the  thalamus.  Within  the  latter,  the  third  neurone  takes 
origin  and  ends  within  the  cerebral  cortex.  The  sensory  end-nuclei  are  also  connected  with 
the  cerebellum,  by  means  of  the  tractus  nucleo-cerebellaris.  Further,  all  of  the  centrally 
coursing  sensory  root-fibres  do  not  terminate  within  the  end-nuclei  of  the  glossopharyngeus 
and  vagus,  since  some  of  them  join  the  descending  tractus  spinalis  of  the  trigeminus  nerve. 

NERVUS   ACCESSORIUS. 

The  spinal  accessory  nerve  presents  a  cerebral  and  a  spinal  portion.  The  fibres  of  the 
cerebral  part  arise  from  a  nucleus,  situated  within  the  caudal  prolongation  of  the  nucleus 
ambiguus  ;  further,  from  a  small  dorsal  nucleus,  which  represents  the  caudal  prolongation  of  the 
dorsal  motor  vagus  nucleus.  The  fibres  of  the  spinal  portion  of  the  accessorius  take  origin 
from  cells  situated  in  the  base  of  the  lateral  horn  and  in  the  dorsolateral  part  of  the  anterior 
horn  of  the  spinal  cord,  as  far  down  as  the  fifth,  or  even  seventh,  cervical  segment  of  the  cord. 

The  path  includes: 

a.  The  central  neurone— horn  cerebral  cortex  to  the  nucleus; 

b.  The  peripheral  neurone — nucleus,   peripheral  nerve,  muscle. 

As  well  known,  the  accessorius  supplies  the  sterno-cleido-mastoideus  and  trapezius 
muscles.  These  fibres  include  the  spinal  portion  of  the  nerve  and  constitute  the  ramus 
externus,  while  the  fibres  of  the  cerebral  portion,  as  the  ramus  internus,  pass  to  the 
vagus,  as  a  part  of  which  they  are  to  be  regarded. 


1 86  THE   FIBRE-TRACTS. 

NERVUS   HYPOGLOSSUS. 

The  nucleus  of  the  hypoglossal  nerve  lies  in  the  floor  of  the  fourth  ventricle,  within  the 
trigonum  nervi  hypoglossi.  The  efferent  fibres  pass  from  the  nucleus,  proceed  ventrally  and 
emerge  from  the  brain-stem  between  the  pyramid  and  the  olivary  eminence.  The  path  includes: 

a.  The  central  neurone — from  cerebral  cortex  (lower  third  of  the  precentral  convolu- 
tion), knee  of  internal  capsule,   nucleus. 

b.  The  peripheral  neurone — nucleus,   peripheral  nerve,   muscle. 

SUMMARY  OF  THE  CHIEF  TRACTS. 

A.    PROJECTION    TRACTS. 

The  entire  sensory  projection  path  from  the  sense-surfaces  (skin,  retina,  labyrinth,  etc. ) 
to  the  sensory  region  of  the  cerebral  cortex,  as  well  as  the  entire  motor  projection  path  from 
the  motor  region  of  the  cerebral  cortex  to  the  muscles,  is  made  up  of  several  paths  of  conduc- 
tion or  projection  systems. 

I.    CENTRIPETAL   TRACTS, 
i.  Ascending  sensory  tracts  from  the  spinal  cord. 

a.  The   path  for   the   conduction  of    impulses   of  touch>  temperature  and  pain  from 
the  trunk  and  the  extremities. 

Neurone  I:  The  impulse  is  conveyed  from  the  periphery  to  the  ganglion -cells 
within  the  spinal  ganglion  and  thence  to  the  spinal  cord  by  the  posterior  roots.  The 
latter  enter  the  spinal  cord  and  end  within  the  gray  substance. 

Neurone  II:  Origin  within  the  gray  substance  of  the  spinal  cord.  The  fibres 
pass,  as  the  axones  of  commissure-cells,  by  way  of  the  anterior  gray  commissure  to  the 
opposite  lateral  column  and  form  the  tractus  spino-thalamicus,  which  higher  up  joins  the 
medial  fillet  and  with  it  ends  within  the  thalamus. 

Neurone  III:  Origin  within  the  thalamus.  Course  to  the  cerebral  cortex,  in  part 
direct  by  way  of  the  internal  capsule,  and  in  part  after  traversing  the  lenticular  nucleus. 
Cortical  ending  within  the  area  of  somatic  sensibility. 

The  conduction  of  impulses  of  contact  or  tactile  sensibility  is  not  limited  to  the 
spino-thalamic  tract,  but  takes  place  also  through  the  long  tracts  of  the  posterior  columns. 

b.  The  path  for  the  conduction  of  the  muscle-sense  from  the  trunk  and  the  extremities. 
Neurone  I:     The  impulse  is  conveyed,  as  in  the  case  of  those  of  touch,  temperature  and 

pain,  first  to  the  spinal  cord.  The  fibres  enter  as  posterior  roots,  do  not,  however,  end  within 
the  gray  substance  of  the  spinal  cord,  but  ascend  within  the  posterior  column  to  the  medulla 
oblongata,  where  they  first  find  their  ending  within  the  nucleus  gracilis  and  cuneatus. 

Neurone  II:  Origin  within  the  posterior  column  nuclei;  course,  after  decussation, 
as  medial  fillet  to  thalamus  and  there  end. 

Neurone  III:     Origin  within  the  thalamus.     Course  to  cortical  somatic  sensory  area. 

The  conduction  of  muscle-sense  is  not  only  by  way  of  the  posterior  column  nuclei  and  the 
medial  fillet,  but  also  shares  the  tracts  passing  to  the  cortex  by  way  of  the  cerebellum,  that  is  by 


CENTRIPETAL  PATHS. 


187 


the  tractus  spino-cerebellaris  ventralis  et  dorsalis.  From  the  cerebellum,  the  conduction  passes 
through  the  superior  cerebellar  peduncle  to  the  thalamus  and  thence  to  the  cerebral  cortex. 
It  is  to  be  noted,  that  tracts  lead  to  the  cerebellum  also  from  the  posterior  column  nuclei. 

The  partly  uncrossed  (muscle-sense  or  deep  sensibility)  and  partly  crossed  (pain 
and  temperature)  conduction  of  sensibility  within  the  spinal  cord,  explains  the  peculiar 
disturbances  of  sensibility  in  hemilesions  of  the  cord,  as  manifested  in  the  Brown-Se"quard 
symptom-complex.  In  hemilateral  lesions  of  the  spinal  cord,  we  find,  on  the  same  side 


Tractus  spino-thalamicus 


Nnc.  grac.  et  cuneat. 


FIG.  172. — The  sensory  tract. 

as  the  lesion,  paralysis  in  consequence  of  interruption  of  the  descending  motor  paths, 
and  disturbances  of  the  deep  sensibility  or  muscle-sense  in  consequence  of  the  involvement 
of  the  ascending  paths  of  the  posterior  column  and  of  the  spino-cerebellar  tracts  ;  while, 
on  the  side  opposite  the  lesion,  are  found  disturbances  of  superficial  sensibility,  pain  and 
temperature  impulses  in  consequence  of  the  crossed  path  of  the  tractus  spino-thalamicus. 
Further,  the  fact  that  the  conduction  of  sensibility,  especially  of  muscle-sense,  also 
takes  place  by  way  of  the  cerebellum,  supplies  the  explanation  of  those  pathological  dis- 
turbances, which  we  designate  as  ataxia  or  errors  of  coordination,  since  the  impulses 
proceeding  from  the  muscles  and  articulations  are  no  longer  transmitted  to  the  cerebellum 
in  consequence  of  the  lesion  of  the  posterior  column  tracts. 


188 


THE   FIBRE-TRACTS. 


2.  Sensory  Tracts  of  the  Cerebral  Nerves. 

a.  The  path  for  the  impulses  of  touch,   temperature  and  pain  from  the    integument 
of  the  head  (with  the  exception  of  the  occipital   region  and  certain  areas  of  the  external 
ear — supplied  respectively  by  the  occipital  and   great  auricular  nerves),  further,  from  the 
conjunctiva,  the  mucous  membranes  of  the  nasal  fossae,  of  the  mouth  and  tongue,  of  the 
palate,  of  the  pharynx,  etc.,  lies  in  the  trigeminus,   the  glossopharyngeus,   or  the  vagus. 

b.  The  path  for  the   impulses  of  orientation  and  movement — muscle-sense— from  the 
face  lies  probably  in  the  trigeminus;  that  from  the  larynx  probably  in  the  vagus. 


—~~Pcsterior  column  tracts 


\    \ 


Trnciut  spino- 
thalanticus 


.^^Tractiis  spino-cercbellai 
dor  sails  et  venlralis 


Reflex  pat 


FIG.  173. — Ascending  tracts  from  the  spinal  cord,     a  4-  6  =  conduction   of    muscle-sense;    c  =  conduction  of  impulses  o» 
pain  and  temperature;    a  +  c  —  conduction  of  tactile  sensibility. 

The  impulse  is  carried  from  the  periphery  to  the  ganglion  of  the  corresponding 
nerve,  and  thence  To  the  end-nucleus  within  the  brain-stem.  To  the  peripheral  neurone  I 
is  added  the  central  neurone'  II.  The  latter  arises  within  the  sensory  end-nucleus,  its 
axone,  the  efferent  nerve-fibre,  passes  upward  with  the  medial  fillet  and  ends  within  the 
thalamus.  From  here  the  neurone  III  extends  to  the  cerebral  cortex. 

c.  The     path     for     the    visceral    impulses,    from     the     lungs,     heart,     oesophagus, 
stomach,  etc.,  lies  in  the  vagus  and  the  sympathetic. 

d.  The    path    for    the   equilibrium    impulses  lies    in    the   vestibular    nerve,    supple- 
mented by  spinal  fibres.     The  path  leads  to  the  cerebellum,  thence  by  way  of  the  superior 
cerebellar  peduncle  to  the  nucleus  ruber  and  the  thalamus,  and  then  to  the  cerebral  cortex. 


CENTRIPETAL    PATHS. 


189 


e.  The  path  for  the  taste  impulses  lies  in  the  glossopharyngeus,  the  intermedius 
and  the  third  division  of  the  trigeminus.  Neurone  I  leads  from  the  periphery 
(the  tongue)  to  the  end-nucleus  (nucleus  of  the  tractus  solitarius);  neurone  II  from 
the  end-nucleus  to  the  thalamus  ;  neurone  III  from  the  thalamus  to  the  cortical  gus- 
tatory centre. 

Concerning  the  paths,  which  serve  to  conduct  the  gustatory  impulses,  the  following 
may  be  noted.  It  is  generally  accepted,  that  the  taste  impulses  from  the  anterior  two- 


Post.  column  tracts 


Pyramidal  tract 


Tract,  spi. 


f: 

'.  spino-thalanticus   '  "      1 


FIG.  174. — Diagram  explaining  Brown-Sequard's  hemilateral  lesion. 

thirds  of  the  tongue  are  conveyed  centrally  by  the  lingual  branch  of  the  trigeminus;  from 
the  posterior  third  of  the  tongue,  by  the  glossopharyngeus.  While  the  course  of  the 
taste-fibres  by  means  of  the  glossopharyngeal  nerve  is  readily  understood,  opinions  con- 
cerning the  path  followed  by  the  taste-fibres  from  the  anterior  two-thirds  of  the  tongue 
vary.  Thus,  it  is  assumed  by  some,  that  these  fibres  run  backward  in  the  chorda  tym- 
pani  to  the  ganglion  geniculi  and  thence  proceed,  either  through  the  great  superficial 
petrosal  nerve  to  the  spheno-palatine  ganglion  and  on  centrally  by  the  maxillary  division 


190  THE   FIBRE-TRACTS. 

of  the  trigeminus,  or  through  the  small  superficial  petrosal  nerve  to  the  otic  ganglion  and 
centrally  by  the  mandibular  nerve.  According  to  •  others,  the  chorda  fibres  reach  the 
glossopharyngeus  by  way  of  the  small  superficial  petrosal  and  the  tympanic  nerve.  It  is 
assumed,  further,  that  not  only  the  chorda  fibres,  but  also  the  gustatory  fibres  of  the 
glossopharyngeus,  by  way  of  the  small  superficial  petrosal  nerve,  reach  the  trigeminus  and 
in  it  pass  centrally.  Finally,  according  to  the  view  which  seems,  perhaps,  the  most 
reasonable,  the  chorda  fibres  pass,  by  way  of  the  chorda  tympani,  to  the  geniculate  gan- 
glion and  thence,  by  way  of  the  nervus  intermedius,  to  the  medulla  oblongata,  where  they 
form  a  descending  root,  which  ends  in  the  nucleus  tractus  solitarii,  that  is,  the  sensory 


FIG.  175. — Possible  conduction  paths  for  gustatory  impulses. 


end-nucleus  of  the  glossopharyngeus.  This  view,  further,  is  supported  by  the  experi- 
mental investigations,  which  have  shown,  that  removal  of  the  Gasserian  ganglion  or  intra- 
cranial  section  of  the  maxillary  and  the  mandibular  divisions  of  the  trigeminus  are  not 
followed  by  loss  of  taste  in  the  anterior  two -thirds  of  the  tongue. 

f.  The    path   for  olfactory  impulses  leads  from    the  olfactory  mucous   membrane  by 
way  of   the  fila   olfactoria   to  the    bulbus   olfactorius,  thence    to    the   primary  centres  and 
from  the  latter  to  the  secondary  or  cortical  olfactory  centre  within  the  gyrus  hippocampi. 

g.  The   path  for    auditory  impulses   lies    within    the    nervus    cochleae.       Neurone    I 
conveys  the   stimulus  from    the  hair-cells  of   Cord's  organ  to  the  end- nucleus.      Neurone 
II  passes  from  the   end-nucleus    to    the  corpus   geniculatum    mediale   and   to    the  inferioi 
colliculus,  the  fibres  forming  the  lateral  fillet.     Neurone  III  unites  the  corpus  geniculatum 
mediale  with  the  cortical  auditory  centre  within  the  gyrus  temporalis  superior. 


CENTRIFUGAL    PATHS. 


191 


h.  The  path  of  the  visual  impulses  lies  within  the  nervus  opticus.  Neurone  I 
extends  from  the  retina  to  the  corpus  geniculatum  laterale,  to  the  pulvinar  and  to  the 
superior  colliculus.  Neurone  II  connects  the  corpus  geniculatum  laterale  and  the  pulvi- 
nar with  the  secondary  or  visual  centre  within  the  cortex  of  the  cuneus. 

II.    CENTRIFUGAL   TRACTS. 

i.  The  efferent  cortico-muscular  or  motor  tract  takes  its  origin  in  the  motor 
region  of  the  cerebral  cortex. 

Neurone  I:  through  the  internal  capsule  (knee  and  anterior  two-thirds  of  the  poste- 
rior limb),  the  basis  or  crusta  of  the  cerebral  peduncle  (middle  three-fifths),  the  pons  and  the 
medulla  oblongata. 


Tractus  rubro-spinalh 
—  Mono-how  — 


Pyramidal  tract 


Cortico-bulbar  tract  to  motor 
cerebral  nerves 


Motor  cerebral  nerves 

Ant.  cerebro-splnal  tract 
Lateral  cerebro-spinal  tract 


FIG.  176. — Motor  paths. 

a.  as  the  tract  of  the  motor  cerebral  nerves,  to  the  contralateral  nuclei  of  the  motor 
cerebral  nerves. 

b.  as  the  pyramidal  tract  proper,  to  the   spinal  cord — crossed  as  the  lateral  pyram- 
idal tract,   uncrossed    (in  the  medulla  oblongata)  as  the  anterior  pyramidal  tract — to   end 
around  the  ventral  horn-cells. 

Neurone  II:  origin  within  the  nuclei  of  the  motor  cerebral  nerves,  peripheral  course 
as  the  motor  cranial  nerves  to  the  muscles ;  or,  in  like  manner,  as 

Neurone  II:  origin  in  the  cells  of  anterior  horn  of  the  spinal  cord,  peripheral 
course  through  the  ventral  roots,  as  the  motor  spinal  nerves  to  the  muscles. 


IQ2 


THE   FIBRE-TRACTS. 


2.  A  special  motor  speech-tract  does  not  exist.      The   speech-tract   is    identical 
with  those  paths,  which,  as  part  of  the  cortico-bulbar  tract,  pass  from  the  cortical  centre 
for  the  facial  and  hypoglossal  nerves  to  the  nuclei  of  the  nerves  necessary  for  speech. 

3.  A  cortico-rubral  motor  path  goes    to  the  spinal  cord  by  way  of   the  nucleus 
ruber,  as  follows  : 

Neurone  I:    from  the  cerebral  cortex  to  the  nucleus  ruber  ; 
Neurone  II:    nucleus  ruber,   tractus  rubro-spinalis,  spinal  cord  ; 
Neurone  HI:    spinal  cord,  anterior  root,   muscle. 

4.  An  indirect  motor  path  passes  to  the  spinal  cord  by  way  of  the  pons  and  the 
cerebellum,  as  follows  :    frontal  and  occipito-temporal  pontile  tract — pontile  nucleus — cere- 
bellar  cortex — nucleus   dentatus    cerebelli — superior    cerebellar    peduncle — nucleus    ruber— 
tractus  rubro-spinalis — spinal  cord — muscle  (Fig.    147). 

5.  In  addition  to  the  above  direct  and  indirect  motor  paths,  others  arise  within  the 
lower  brain-centres  and  pass  spinalward  ;    such  are  the  tractus  rubro-spinalis,   the  tractus 
tecto-spinalis  and  the  tractus  vestibulo-spinalis. 

B.    REFLEX    TRACTS. 

The  simplest  reflex  path  is  established  by  the  reflex  collaterals.  In  this  case 
only  two  neurones  share  the  entire  path,  the  transference  from  the  centripetal  to  the 
centrifugal  neurone  being  accomplished  by  means  of  the  collaterals  given  off  directly 

from  the  centripetal  or  afferent  neurone. 
The  release  of  the  reflex  may  be  in- 
duced, however,  by  intercalated  neurones. 
Thus,  between  the  centripetal  and  the  cen- 
trifugal neurone  a  third  neurone  may  inter- 
vene, thereby  making  possible  the  transfer- 
ence of  the  impulse  conveyed  by  a  single 
centripetal  neurone  to  several  centrifugal 
ones.  .  Such  intercalated  neurones,  for  ex- 
ample, are  the  association-cells  of  the  spinal 
cord,  which  distribute,  by  means  of  their 
axones  and  collaterals,  impulses  to  many 
cells  within  the  cord-segments  of  higher  and 
lower  levels.  To  this  category  belongs,  fur- 
ther, the  posterior  longitudinal  bundle.  Im- 
pulses carried  to  Deiters'  nucleus  by  the 
vestibular  nerve  may  be  distributed  to  the 
nuclei  of  the  eye-muscles  and  to  the  motor 
cells  of  the  cord  by  means  of  fibres,  which 
proceed  from  Deiters'  nucleus  and  run 
within  the  posterior  longitudinal  bundle. 
In  consequence  of  the  introduction  of  sev- 
eral neurones  between  the  centripetal  and 
the  centrifugal  conduction,  the  entire  reflex 

FIG.  177- — Reflex  paths  in  the  spinal  cord.     Broken  lines  rep- 
resent neurones  distributing  impulses  to  other  levels.  mechanism     may    become    very    complex. 


REFLEX    PATHS. 


193 


The  cerebellum,  with  its  afferent  and  efferent  paths,  calls  for  special  consideration. 
The  cerebellum  is  the  centre  for  the  reflex  and  unconscious  maintenance  of  equilibrium, 
during  rest  as  well  as  during"  changes  in  the  position  of  the  centre  of  gravity.  The 
centripetal  paths  lie  especially  within  the  nervus  vestibuli  and  within  the  ascending  fibre- 
systems  from  the  spinal  cord  and  from  the  medulla  oblongata.  These  ascending  paths 
from  the  cord  are  the  tractus  spino-cerebellaris  dorsalis  and  ventralis  ;  from  the  medulla 


Fuse,  longitudinalis 
medlalis 


Tract,  vestilndo-spinalis  . 
Posterior  column  tract.  - 


Tract,  spino-reretellar 
ventral  is  (Cowers) 


Tract.  sf>ino-cerebellaris 
dorsalis  (Flechsig) 


X 


FIG.   178. — Spino-cerebellar  and  cerebellofugal  tracts.     Vestibulo-cerebellar  tract-system  of  Deiters"  nucleus. 

oblongata  the  fibres  arising  within  the  nucleus  gracilis  and  cuneatus.  An  indirect  con- 
duction from  the  spinal  cord  is  perhaps  effected  by  the  tractus  spino-olivaris  or  Helweg's 
triangular  tract,  which  ends  within  the  inferior -olivary  nucleus,  and  thence  by  the  tractus 
olivo-cerebellaris  to  the  cerebellum,  by  way  of  the  restiform  body.  The  direct  and  the 
indirect  sensory  cerebellar  tract,  as  well  as  the  tracts  from  the  quadrigeminal  region,  are 
also  included  among  the  centripetal  paths.  By  means  of  the  cerebellofugal  tracts,  impulses 
may  be  carried  from  the  cerebellum  to  other  paths  and  by  means  of  the  latter,  in  turn, 
be  transferred  to  motor  paths.  The  chief  cerebellofugal  tracts  proceed  from  Deiters' 
nucleus  and  from  the  nucleus  dentatus.  From  Deiters'  nucleus  arise  the  tractus  vestibule- 


i94 


THE   FIBRE-TRACTS. 


spinalis  and  the  posterior  longitudinal  bundle,  the  last-named  system  coming  into  relation 
with  the  spinal  cord  and  with  the  nuclei  of   the  nerves  supplying  the  ocular  muscles — 


Tract  from  pans  to 
cerebellum  (middle  peduncle) 


Tracts  oj  post,  column  _ 


Tract,  vestibule  spinalis 


Tract,  rubro-spinalis 


Tract,  spino-cerebellaris 


FIG.  179. — Cerebellopetal  and  cerebellofugal  tracts. 


that  is,  binding  together  the  centres  concerned  in  maintaining  equilibrium  and  relations 
to  space.  From  the  nucleus  dentatus  arises  the  superior  cerebellar  peduncle,  whose  fibres 
end  within  the  nucleus  ruber,  whence  the  tractus  rubro-spinalis  passes  to  the  spinal  cord. 


REFLEX    PATHS. 


195 


It  is  to  be  noted,  that  the  relations  of  the  cerebellar  hemispheres  with  the  spinal  cord 
are  homolateral  or  of  the  same  side.  Additional  cerebellofugal  paths  are  those  which,  as 
the  tractus  tegmentalis  pontis  et  bulbi,  run  within  the  tegmental  region  of  the  pons  and 
medulla  oblongata,  whereby  transference  to  motor  nuclei  may  in  turn  be  effected. 

If   the  maintenance  of   equilibrium  be  adjusted  to  a  voluntary  movement,  the  cere- 
bellum is  also  directly  stimulated  from  the  cerebral  cortex.     The  paths  for  such  impulses 


Tracts  coming  from  higher  respiratory 
centres  to  nucleus   respiratorius 


tral  (nitcleo-cortical) 
tract  of  vagus 


Sensory  ena-nucleus 
of  -vagus 


Sanguis 


Plex.  brachial. ' 


Nn.l-horacales 


FIG.  180. — Schematic  representation  of  the  tracts  chiefly  concerned  in  respiration. 

are  the  frontal  and  the  temporo-occipital  cortico-pontile  tracts,  which  end  in  the  pontile 
nuclei,  whence  the  conduction  to  the  cerebellum  is  by  the  middle  cerebellar  peduncle. 
In  addition,  the  pontile  nuclei  are  under  the  influence  of  the  pyramidal  tract,  from  which, 
within  the  pons,  collaterals  are  given  off  to  the  nuclei.  By  means  of  the  superior  cerebellar 
peduncle  (cerebellum-nucleus  ruber — thalamus — cortex),  the  cerebellum  sends  impulses  to 
the  cerebral  cortex  and  thereby  influences  conscious  innervation  (Figs.  145  and  147). 

Besides    the    cerebellum,    other    organs    that    preside    over    reflex    activity    call    for 
mention.     Such  organs,  in  the  first  place,  are  the  thalamus  and  the  corpora  quadrigemina. 


i96  THE   FIBRE-TRACTS. 

The  centripetal  paths  of  the  thalamus  are  :  the  ascending  tract  of  the  medial  fillet,  the 
fibres  of  the  optic  tract  ending  within  the  pulvinar,  the  fibres  from  the  olfactory  centres 
and  the  fibres  from  the  cerebellum  by  way  of  the  superior  peduncle.  The  thalamofugal 
paths  lie  within  the  tractus  thalamo-spinalis,  the  tractus  rubro-spinalis  and  the  central 
tegmental  tract.  By  means  of  the  connections  between  the  thalamus  and  the  cerebral 
cortex,  impulses  coming  from  the  periphery  are  carried  to  the  cortex  and,  in  reversed 
direction,  activities  occurring  withjri  the  cerebrum  are  transferred  to  lower  lying  centres. 

The  centripetal  path  of  the  superior  colliculus  lies  within  the  tractus  opticus  and 
partly  within  the  lateral  fillet ;  that  of  the  inferior  colliculus  within  the  lateral  fillet.  A 
centripetal  path  of  the  corpora  quadrigemina  is  afforded  also  by  the  ascending  tractus 
spino-tectalis,  associated  with  the  tractus  spino-thalamicus.  Fibres  pass  from  the  quadri- 
geminal  region  to  the  cerebellum  and  an  important  descending  path  forms  the  tractus 
tecto-spinalis,  the  path  from  the  quadrigeminal  bodies  to  the  spinal  cord.  Since  fibres 
from  the  optic  and  acoustic  nerves  end  within  the  quadrigeminal  region  and  the  path 
effects  the  transference  of  impulses  of  these  nerves  to  the  spinal  cord,  the  tecto-spinal 
tract  is  also  known  as  the  msuo-auditory  reflex  path. 

The  foregoing  by  no  means  completes  the  enumeration  of  the  reflex  paths,  since 
throughout  the  brain-stem  course  numerous  additional  tracts,  which  serve  to  unite  func- 
tionally related  centres.  In  this  connection,  it  is  only  necessary  to  recall  the  complex 
mechanism  of  the  medulla  oblongata,  in  which  different  nuclei  are  brought  into  the  most 
varied  relations,  whereby  numerous  simple,  as  well  as  the  most  complex,  reflex  proc- 
esses are  effected.  While  it  is  impracticable  here  to  consider  all  such  reflex  paths,  in 
order  to  obtain  some  notion  of  such  complicated  mechanisms,  we  may  represent,  by 
means  of  a  simple  diagram,  the  centres  and  tracts  chiefly  concerned  in  respiration. 

Respiration  is  maintained  by  the  stimulus  carried  to  the  respiratory  centre .  through 
the  circulation.  In  addition,  the  reflexes  transmitted  by  the  vagus  also  come  into  con- 
sideration. In  Fig.  1 80,  the  respiratory  centre  is  represented  by  the  nucleus  respiratoriiis 
within  the  formatio  reticularis.  This  nucleus  stands  in  close  relation  with  the  sensory 
end-nucleus  of  the  vagus,  since  impulses  are  conveyed  to  it  by  the  collaterals  given  off 
from  the  central  vagus-tract.  Moreover,  as  indicated  in  the  figure,  the  nucleus  respira- 
torius  is  also  under  the  influence  of  the  higher  lying  respiratory  centres.  By  means  of 
the  paths  passing  from  the  respiratory  nucleus,  as  well  as  by  the  farther  connecting 
neurones,  the  irrpulse  is  transferred  to  the  motor  nuclei  of  certain  cerebral  nerves  and 
the  gray  substance  of  the  spinal  cord  and,  thence,  is  carried  by  the  motor  fibres  to  the 
muscles  concerned  in  respiration.  Thus,  the  impulse  is  carried  by  the  phrenic  nerve  to 
the  diaphragm;  by  the  thoracic  nerves  to  the  intercostales  and  levatores  costarum;  by  the 
cervical  plexus  to  the  scalene,  sterno-hyoid  and  sterno-thyroid  muscles  (depression  of  the 
larynx);  by  the  brachial  plexus  to  the  rhomboidei  and  pectoralis  minor;  by  the  accessorius 
to  the  sterno-cleido-mastoid  and  trapezius;  by  the  vagus  to  the  crico-arytaenoideus  posticus 
and  thyero-arytaenoideus  (widening  of  the  vocal  cleft)  and  the  levatores  veli  palatini  et 
uvulae  (elevation  of  the  soft  palate  and  the  uvula);  the  facial  nerve  to  the  facial  muscles 
(widening  of  the  nasal  apertures  and  the  oral  cavity).  The  paths  passing  from  the  nucleus 
respiratorius  course  in  the  medulla  oblongata  within  the  formatio  reticularis,  and,  as  shown 
in  the  accompanying  diagram,  numerous  motor  nuclei  are  brought  into  common  activity 
by  means  of  these  association  tracts  (Fig.  220). 


ASSOCIATION    PATHS.  197 


C.    ASSOCIATION    TRACTS. 

.  When  discussing  cerebral  localization,  it  was  pointed  out,  that  the  various  divisions 
of  the  brain  were  divided  in  a  general  way,  according  to  their  function,  into  higher  and 
lower  parts.  Functionally  the  highest  division  is  the  cerebrum,  with  the  cerebral  cortex; 
the  lower  divisions  intervene  between  the  spinal  cord  and  the  cerebrum  and  include  the 
medulla  oblongata,  the  pons  and  the  cerebellum,  the  mid-brain  and  the  diencephalon  or 
inter-brain. 

All  nerve  tracts,  which  convey  to  the  central  nervous  system  the  most  varied 
impulses  from  the  individual  sense-organs  and  the  various  organs  within  the  body,  find 
their  immediate  ending  within  the  lower  brain-centres;  within  these  lower  centres  arise 
efferent  paths,  by  means  of  which  the  stimuli  received  are  again  projected  towards  the 
periphery  and  transferred  to  the  organs  of  movement.  In  this  manner  are  brought  about 
all  those  movements  that  we  designate  as  simple  and  complex  reflexes,  which  occur 
without  participation  of  our  consciousness.  The  impulses  conveyed  to  the  central  nervous 
system,  however,  are  not  confined  to  the  subcortical  centres,  but  are  carried  by  other 
paths  to  the  cerebral  cortex,  where,  in  the  appropriate  sensory,  centres,  impulses  are 
called  forth  which  psychically  correspond  to  what  we  designate  as  sensation.  This  impulse 
within  the  sensory  cortical  centres  continues,  so  long  as  the  stimulus  continues.  With 
the  stimulus,  the  impulse  disappears  and  therewith  the  sensation  also  ceases.  We  are 
able,  however,  to  picture  an  object,  even  when  we  no  longer  perceive  it,  or  to  recognize 
it  when  it  again  appears.  Therefore,  on  its  first  appearance,  the  stimulus  must  have 
called  forth  a  permanent  impulse,  in  addition  to  the  vanishing  sensory  impulse;  the  latter 
is  designated  the  concept  impulse.  The  retention  of  this  impulse  makes  possible  the 
recognition,  the  proving,  or  the  representation  of  the  object;  that  is,  there  remain  per- 
sistent traces  of  previous  sensory  or  motor  impulses,  the  so-called  latent  dispositions. 
These  latent  dispositions  or  subconscious  impressions,  when  later  awakened  by  new  impulses, 
render  possible  the  conscious  memory  or  conception  of  sensation  and  movement.  The 
ability  to  call  into  activity  and  to  convert  the  latent  dispositions  or  impressions  into 
conceptions  is  what  we  call  thought. 

In  addition  to  this  mnemonic  function,  the  cerebrum  possesses  the  associative  function. 
One  conception  can  awaken  others  by  reason  of  the  linking  together  of  the  latent  dis- 
positions. By  union  of  partial  conceptions  (visual,  gustatory,  olfactory,  tactile  and  other 
sensations),  the  complete  conception  is  attained;  by  the  blending  of  the  complete  con- 
ceptions, the  general  conceptions  are  formed.  In  this  way  are  "reproduced"  entire 
complexes  of  conceptions,  which  are  definitely  connected  and,  as  it  were,  lie  prepared; 
it  may  be,  however,  that  certain  complexes  of  conception  are  arranged  in  other  and  new 
sequences,  new  conceptions  being  thereby  "produced."  The  associative  function  consists, 
therefore,  in  the  reproduction  and  production  of  conceptions,  and  on  this  possibility  of  a 
definite  sequence  of  conceptions  depends  the  exercise  of  the  higher  psychic  processes, 
that  is,  thought. 

By  means  of  these  associative  processes,  the  individual  cortical  areas  within  the 
same  projection  and  memory  fields,  as  well  as  the  different  projection  and  memory  fields, 
are  brought  into  connection  with  one  another.  Such  connection  between  the  dispositions 


PIG.  181. — Schematic  representation  of    the  physiologically  different  conductions.     Red,  centrifugal   tracts;    blue,  centrip- 
etal tracts;  black,  intercentral  tracts. 


ASSOCIATION   PATHS.  199 

or  residues  of  the  same  kind  exists  everywhere  within  the  corresponding  cortical  areas. 
The  association  between  residues  of  different  kinds,  as  well  as  the  connection  of  projection 
areas  with  memory  centres  and  of  the  various  projection  and  memory  centres  with  one 
another,  is  established  by  means  of  the  association  fibres,  which  as  short  and  long  fibres 
unite  adjoining  convolutions  and  remote  regions  respectively. 

Since,  however,  the  widely  different  processes  of  the  outer  world  and  of  the  body 
proper  give  rise  to  the  formation  of  manifold  impressions  and  to  the  exercise  of  the 
most  simple  as  well  as  the  highest  psychic  processes,  something  further  always  occurs. 
The  influences  taken  up  by  the  organism  react  outwardly,  since  they  always  find  expres- 
sion in  the  various  movements  of  the  organs.  While  the  purely  reflex  reactions  are 
carried  out  unconsciously,  through  the  agency  of  the  lower  brain-centres  and  without 
the  participation  of  the  cerebrum,  the  voluntary  movements,  our  conduct  and  voluntary 
acts  are  dependent  upon  the  activity  of  the  cerebral  cortex,  every  action,  indeed,  being 
determined  by  conceptions  and,  in  the  final  analysis,  by  kinaesthetic  or  motor  concepts. 
These  relations  will  be  best  understood,  if,  in  conclusion,  we  consider  more  closely  those 
most  important  movements  concerned  in  speech,  by  means  of  which  our  entire  sensations, 
conceptions  and  thoughts  find  expression. 

When  discussing  cerebral  localization,  it  was  pointed  out,  that  in  right-handed, 
therefore,  in  the  majority  of  individuals,  the  speech-zone,  with  its  different  centres,  was 
located  within  the  left  hemisphere.  The  chief  centres  include  (Fig.  182):  the  sensory 
speech-centre  (A),  within  the  posterior  third  of  the  superior  temporal  convolution,  where 
the  memory-pictures  of  the  heard  words  are  deposited — the  centre,  therefore,  for  the 
memory  of  word-tones — and  the  motor  speech-centre  (M),  within  the  posterior  third  of 
the  inferior  frontal  convolution,  in  whose  cells  the  memory-pictures  of  the  spoken  words 
lie  and  on  whose  integrity  depends  the  ability  to  carry  out  the  coordinated  movements 
of  certain  muscles  necessary  for  speech. 

These  two  speech-centres,  the  sensory  and  the  motor,  stand  in  close  relation  with 
each  other,  the  latter  dependent  upon  the  former,  since  speech  is  acquired  by  repetition 
of  the  word-sounds  heard.  On  observing  the  development  of  speech  in  the  child,  we  find 
in  the  connection  of  these  two  centres  the  basis  for  the  possibility  of  pronouncing  by 
repeating  but  without  understanding.  The  development  of  speech  teaches,  moreover,  that 
speech  proper,  that  is,  the  intelligent  utterance  of  sounds,  in  contrast  to  their  mere  repeat- 
ing, is  preceded  by  an  understanding  of  speech  without  speaking — a  stage  of  ' '  normal 
deaf-mutism."  The  child  understands  much,  but  speaks  little  or  nothing  of  what  it 
understands  ;  it  is,  for  the  time,  deaf-mute.  Therefore,  an  intimate  connection  is  early 
established  between  the  memory  of  the  word-sound  or  the  acoustic  word  (A)  and  the 
idea  (-5).  In  Fig.  182,  this  close  connection,  as  well  as  that  between  the  sensory  and 
motor  speech-centres,  is  represented  by  the  double  line,  A  —  B.  In  this  relation  it 
should  be  emphasized,  that  the  idea-centre  (Z?)  is  represented  as  a  definitely  bounded 
cortical  area  only  as  a  schematic  expedient,  and  that,  as  a  matter  of  fact,  we  must  con- 
ceive the  formation  of  the  idea  as  a  complex  process  involving,  more  or  less,  the  entire 
cerebral  cortex. 

From  this  speech-comprehension  without  speaking  (a1  —  a2  —  a  —  A  —  B}  and  the 
first  mere  repeating  of  spoken  words  (a1  —  a2  —  a  —  A  —  M  —m  —  m1  —  m2),  later  comes 
the  repeating  of  words  with  the  understanding  of  speech;  that  is,  speech  proper.  The 


THE   FIBRE-TRACTS. 


x    M     v 


Tkojtti 


Speech 


Writing 


Fig.  182. — Scheme  of  spoken  and  written  speech.  B,  conception  centre;  M,  motor  speech-centre  (Broca);  A,  sensory 
speech-centre  (Wernicke);  O,  visual  letter-centre;  m,  motor  centre  (facial,  lingual  and  laryngeal  musculature);  a,  auditory 
centre;  o,  visual  centre:  H,  motor  centre  for  hand;  m1  m*  hl  h*  —  cortico-muscular  tracts  for  speaking  and  writing; 
<ji  o2  o1  o2  —  auditory  and  visual  tracts. 


ASSOCIATION   PATHS.  201 

latter  first  takes  the  path  :  B  —  A  —  M—  m  —  m1  —  m2 ;  later,  in  consequence  of  the 
connection  B  —  M,  it  follows  :  B  —  M—  m  —  m1  —  m2.  The  centre,  m,  represents  the 
motor  centre  proper,  in  the  lower  third  of  the  precentral  convolution  (the  motor  centre 
for  the  face,  tongue,  and  larynx).  The  path,  m1,  :is  the  motor  cortico-bulbar  tract,  which 
passes  through  the  knee  of  the  internal  capsule,  the  crusta  or  base  of  the  cerebral 
peduncle  to  the  nuclei  of  the  appropriate  motor  cerebral  nerves.  Path  m2  is  the  periph- 


Motor  hand*entre 


Motor  speech- 
centre  (Broca) 


ng-centr, 


FIG.    183. — Connections    of  the  individual   centres   of  the   speech   zone,   represented   on   a  horizontal   section 

through   the   brain. 

eral  motor  neurone  from  the  motor  nucleus  to  the  muscle.  Likewise,  close  to  the 
sensory  speech-centre  (A),  the  auditory  centre  proper  is  represented  (a).  The  path  a1 
shows  the  course  of  the  auditory  tract  as  far  as  the  medial  geniculate  body  ;  the  path  a2 
is  the  last  neurone  in  the  auditory  tract,  which  extends,  by  way  of  the  internal  capsule, 
from  the  geniculate  body  to  the  cortical  auditory  centre. 

The  foregoing  connections  represent  speech  in  the  more  limited  sense;  later,  as  the 
result  of  learning  written  speech — reading  and  writing — the  expansion  to  speech  in  its 
widest  sense  follows.  By  written  speech  is  understood  the  speech  of  the  letters  ;  the  latter 
are  to  be  regarded  not  as  signs  for  ideas,  as  hieroglyphics,  but  as  signs  for  sounds.  We 


202  THE   FIBRE-TRACTS. 

learn  to  separate  the  individual  words  into  syllables  and  letters,  each  simple  sound,  vocals 
and  consonants,  being  associated  with  a  visual  letter-picture ;  by  copying  the  optical 
picture  of  the  letter  we  learn  to  write.  The  sensory  speech-centre  or  the  acoustic  word 
now  comes,  therefore,  into  closer  relation  with  the  visual  apparatus.  Not  only  the 
acoustic  word,  but  also  the  motor  word,  or  the  centre  for  the  motor  memory-pictures  of 
the  words,  becomes  connected  with  the  visual  letter-centre,  or  visual  centre,  O,  within 
the  gyrus  angularis ;  here  the  memory-pictures  of  the  written  characters  are  deposited, 
since  for  reading  the  sensory  and  motor  speech-centres  are  necessary.  For  writing,  more- 
over, connection  is  established  between  the  visual  centre,  O,  and  the  motor  centre  for 
the  upper  extremity  within  the  middle  part  of  the  precentral  convolution',  the  centre,  N, 
for  the  musculature  of  the  hand,  wherein  the  grapho-motor  memories  are  developed 
through  practice.  In  Fig.  182,  this  locality  is  represented  by  two  superimposed  ovals, 
since  the  existence  of  a  distinct  writing-centre  is  not  accepted. 

Reading  is  accomplished,  therefore,  by  the  path:  ol  —  o2  —  o  —  O  —  A  or  M—B; 
spontaneous  writing  by  :  B  —  A  or  M—  O  —  H—  hl  —  h2.  The  path  o1  represents  the 
first  neurone  of  the  visual  path  leading  to  the  lateral  geniculate  body;  the  path  o2  is  the 
second  neurone  from  the  geniculate  body,  by  way  of  the  internal  capsule,  to  the  visual 
centre  proper,  o.  The  latter  is  shown  in  the  diagram  in  the  occipital  pole,  but,  as  well 
known,  the  centre  is  localized  chiefly  within  the  cortex  of  the  cuneus,  particularly  sur- 
rounding the  calcarine  fissure.  The  path,  A1,  represents  the  course  of  the  motor  tract 
from  the  arm  centre  through  the  internal  capsule  and  the  brain-stem  to  the  spinal  cord; 
the  path,  h2>  is  the  peripheral  motor  neurone  to  the  muscles  of  the  hand. 

In  Fig.  183,  the  connections  of  the  individual  speech-centres  are  represented  schem- 
atically in  horizontal  section.  The  course  of  the  tracts  from  one  hemisphere  to  the  other, 
through  the  corpus  callosum,  is  to  be  noted. 

On  Fig.    182,   we  may  trace  the  following  paths  : 

Speech  comprehension  :   a1  —  a2  —  a  —  A  —  B  ; 

Repeating  words  :   a1  —  a2  —  a  —  A  —  M—  m  —  ml  —  m2 : 
Spontaneous  speech  :    B  —  A  —  M—  m  —  m1  —  m* 

B  -  M-  m-ml-  m* 
Reading  :    o1  -  o2  -  o  -  O  -  A  or  M—  B; 
Reading'aloud  :   o1  —  o2  —  o —  O  —  A  or  M—B — M—m  —  m1 — m2; 
Spontaneous  writing  :    B  —  A  or  M  —  O  —  H  —  hl  —  h2; 

Copying  :    o1  —  o2  —  o  —  O  —  H—  hl  —  h2 ; 
Dictated  writing  :    a1  —  a2  —  a  —  A  or  M—  O  —  H  —  hl  —  h2. 

At  the  same  time,  the  diagram  explains  the  different  types  of  the  disturbances  ol 
speech  or  aphasia. 

A  lesion  of  the  speech-centre,  M,  leads  to  cortical  motor  aphasia.  The  patient  can 
neither  speak  spontaneously,  nor  repeat;  moreover,  since  reading  and  writing  depend 
upon  the  integrity  of  the  sensory  as  well  as  of  the  motor  speech-centre,  reading,  spon- 
taneous writing  and  dictated  writing  are  also  impaired.  On  the  other  hand,  the  patient 
understands  what  is  spoken,  since  A  is  intact,  and  can  copy  writing. 


ASSOCIATION   PATHS.  203 

A  lesion  of  the  sensory  speech-centre,  A,  leads  to  cortical  sensory  aphasia.  In  the 
first  place,  comprehension  of  speech  is  lost;  further,  repetition,  reading  and  dictation 
writing  are  suspended,  while  spontaneous  writing  and  copying,  as  well  as  speech,  are 
retained.  In  speaking,  however,  the  patient  manifests  the  symptoms  of  paraphasia,  that 
is,  the  interpolation  of  incorrect  words  and  exchange  and  mutilation  of  words. 

Destruction  of  both  chief  centres,  the  motor  and  the  sensory,  leads  to  total  aphasia. 

When  the  efferent  path  from  the  motor,  speech-centre,  M,  is  interrupted  by  a  sub- 
cortical  effusion,  the  clinical  picture  of  subcortical  motor  aphasia  or  word-dumbness 
appears;  when  the  lesion  involves  the  path  to  the  sensory  speech-centre,  subcortical  sen- 
sory aphasia  follows.  These  subcortical  aphasias  leave  inward  speech  intact,  and  the 
ability  to  read  and  write  are  retained.  On  the  other  hand,  in  subcortical  motor  aphasia, 
voluntary  speech,  repeating  and  reading  aloud,  are  suspended  or  involved;  in  subcortical 
sensory  aphasia,  speech-comprehension,  repeating  and  dictation  writing  are  wanting  or 
impaired. 

If  the  path  from  the  idea-centre  to  the  motor  speech-centre  {BM}  be  interrupted, 
the  patient  is  said  to  be  affected  with  transcortical  motor  aphasia,  with  loss  of  voluntary 
speech  and  writing;  if  the  path  from  the  sensory  centre  to  the  idea-centre  be  broken, 
the  resulting  condition  is  termed  transcortical  sensory  aphasia,  with  loss  of  the  compre- 
hension of  speech  and  of  writing. 

An  interruption  of  the  path  uniting  the  sensory  and  motor  speech-centres  {AM} 
leads  to  the  so-called  conduction  aphasia.  The  ability  of  repeating  words  is  impaired ; 
speech  and  comprehension  of  writing  and  the  ability  to  copy  are  retained,  as  well  as 
spontaneous  speech  and  writing;  the  performance  of  these  functions,  however,  is  attended 
with  the  manifestations  of  paraphasia  and  paragraphia. 


PART  III. 

SERIAL  SECTIONS  THROUGH  THE  BRAIN-STEM 
OF  A  FOUR-YEAR-OLD  CHILD 


FROM  THE  ANTERIOR  END  OF  THE  CORPUS  CALLOSUM  TO  THE 
QUADRIGEMINAL  REGION 


FROM   CORPUS   CALLOSUM  TO  QUADRIGEMINAL  REGION.  207 


208 


SERIAL   SECTIONS   THROUGH   THE   BRAIN-STEM. 


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FROM    CORPUS    CALLOSUM  TO  QUADRIGEMINAL  REGION. 


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SERIAL   SECTIONS   THROUGH   THE   BRAIN-STEM. 


FROM    CORPUS  CALLOSUM  TO  QUADRIGEMINAL  REGION.  211 


212 


SERIAL    SECTIONS   THROUGH   THE   BRAIN-STEM. 


FROM    CORPUS   CALLOSUM  TO  QUADRIGEMINAL  REGION.  213 


214 


SERIAL   SECTIONS   THROUGH   THE   BRAIN-STEM. 


FROM    CORPUS   CALLOSUM  TO  QUADRIGEMINAL  REGION.  215 


216 


SERIAL   SECTIONS   THROUGH   THE   BRAIN-STEM. 


FROM    CORPUS   CALLOSUM  TO  QUADRIGEMINAL  REGION.  217 


218 


SERIAL   SECTIONS   THROUGH   THE   BRAIN-STEM. 


FROM   CORPUS    CALLOSUM  TO  QUADRIGEMINAL  REGION.         219 


SERIAL   SECTIONS   THROUGH   THE   BRAIN-STEM. 


FROM    CORPUS   CALLOSUM  TO  QUADRIGEMINAL  REGION.  221 


SERIAL   SECTIONS   THROUGH    THE   BRAIN-STEM. 


FROM    CORPUS   CALLOSUM  TO  QUADRIGEMINAL  REGION.  223 


SERIAL   SECTIONS   THROUGH   THE   BRAIN-STEM. 


FROM    CORPUS   CALLOSUM  TO  QUADRIGEMINAL  REGION. 


225 


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SERIAL    SECTIONS   THROUGH    THE   BRAIN-STEM. 


FROM    CORPUS   CALLOSUM  TO  QUADRIGEMINAL  REGION. 


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SERIAL   SECTIONS   THROUGH   THE    BRAIN-STEM. 


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FROM   CORPUS   CALLOSUM  TO  QUADR1GEMINAL  REGION. 


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SELECTED    REFERENCES. 

The  following  list  includes  important  books  and  monographs  relating  to  the  anatomy  of 
the  Central  Nervous  System,  which  the  student  may  consult  with  advantage.  Extended  bib- 
liographies will  be  found  in  Obersteiner  and  in  the  year-books  of  Anatomy  and  Neurology. 

BARKER,  L.  F.  The  Nervous  System  and  its  Constituent  Neurones.  1899. 
BECHTEREW,  W.  Die  Leitungsbahnen  im  Gehirn  und  Ruckenmark.  1899. 
BRODMANN,  K.  Die  Cortexgliederung  des  Menschen.  Jour.  f.  Psychol.  u.  Neurol., 

Bd.   X.,  1907. 
BRODMANN,   K.      Die  feinere  Anatomic  des  Grosshirns.      Lewandowsky's    Handbuch  der 

Neurologic,  1910. 

CAJAL,   S.   RAMON  Y.      El   systema    nervioso   del    Hombre  e    de   los  Vertebrados.      1902. 
CHARPY,    A.       Systeme     Nerveux.       In    Poirier's    Traite"    d' Anatomic     Humaine,    Tome 

III.,    1899. 

CUNNINGHAM,   D.   J.     Surface    Anatomy   of   the    Cerebral    Hemispheres.      1892. 
DALTON,  J.   C.     Topographical    Anatomy   of  the    Brain.     Atlas.      1885. 
DEJERINE,  J.     Anatomic  des    Centres    Nerveux.      1901. 

DOGIEL,   A.   S.     Der  Bau  der  Spinalganglien  des  Menschen  und  der  Saugethiere.      1908. 
DONALDSON  AND  DAVIS.     Areas,  of    Cross-Sections    of    the    Spinal    Cord   at    the    Level 

of   each    Spinal    Nerve.     Journal    of   Comparative   Neurology,  Vol.   XIII.,    1903. 

EDINGER,   L.     Vorlesungen   ueber   den    Bau   der  nervosen    Zentralorgane.      1908. 
FLECHSIG,   P.      Die    Leitungsbahnen    im    Gehirn   und    Ruckenmark.      1876. 
GEHUCHTEN,   VAN  A.     Anatomic   du    Systeme    Nerveux   del'Homme.      1906. 
HABERLIN,   C.     Zur  Topographic  der  Hirnventrikel.      Arch.  f.  Anat.   u.   Entwick.      1909. 
HELD,   H.      Die    Enstehung   des    Nervengewebes.      1909. 
His,  W.      Die   Formentwickelung   des    menschlichen   Vorderhirns.      1889. 
His,  W.      Die    Entwickelung   des   menschlichen  Gehirns.      1904. 

HORSLEY,  V.     The    Cerebellum  :      Its    Relation    to    Spacial    Orientation   and    to    Loco- 
motion.     Boyle    Lecture.      1905. 

JACOBSOHN,   L.      Ueber    die     Kerne     des     menschlichen    Hirnstamms.       Akademie     der 
Wissenschaften,    Berlin.      1911. 

JAKOB,   C.     Vom   Tierhirn   zum    Menschenhirn.     Atlas.      1911. 
JOHNSTON,  J.    B.     The   Nervous   System    of   Vertebrates.      1906. 

JOHNSTON,  J.    B.     The    Morphology   of   the    Fore-Brain    Vesicle    in  Vertebrates.     Journal 
of   Comparative    Neurology   and    Psychology.     Vol.    XIX.,    1909. 

279 


28o  SELECTED    REFERENCES. 

KAES,  T.      Die   Grosshirnrinde   des    Menschen    in    ihren    Massen    u.     in    ihrem    Faserge- 

halt.      1907. 

KOELLIKER,   A.      Handbuch   der   Gewebelehre.      Band  II.,  1896. 
KUPFFER,   K.     Die    Morphologic    des    Central  nervensystems.      In    Hertwig's    Handbuch 

der    Entwickelungslehre.      Bd.    II.,    Th.    3  ,    1906. 
LENHOSSEK,  M.      Der  feinere  Bau  des  Nervensystems.      1895. 

LEWANDOWSKY,   M.     Handbuch  der  Neurologic.      I.   Allgemeine  Neurologic.      1910. 
MALONE,   E.     Ueber  die  Kerne  des  menschlichen  Diencephalon.     Akademie  der  Wissen- 

schaften,   Berlin.      1910. 

MARBURG,  O.  Mikroskopisch-topographischer  Atlas  des  menschlichen  Zentralnervensys- 
tems.  1910. 

MONAKOW,  C.  Der  rothe  Kern,  die  Haube  u.  die  Regie  subthalamica  bei  einige  Sauge- 
thieren  u.  bei  Menschen.  Arbeiten  a.  d.  Hirnanatomischen  Institut  in  Zurich. 
Bd.  III.,  1909. 

NEBELTHAU,   E.     Schnitte  durch  das  menschliche  Gehirn.      1898. 

NEUMAYER,  L.  Histo-  und  Morphogenese  des  peripheren  Nervensystems,  der  Spinal- 
ganglien  und  des  Nervus  sympathicus.  In  Hertwig's  Handbuch  der  Entwickel- 
ungslehre. Bd.  II.,  Th.  3,  1906. 

OBERSTEINER,  H.     Anleitung  beim  Studium  des  Baues  der  nervosen  Centralorgane.      1912. 

PARKER,  G.  H.  The  Phylogenetic  Origin  of  the  Nervous  System.  Anatom.  Record, 
Vol.  IV.,  1910. 

RETZIUS,   G.     Das  Menschenhirn.      1896.      Atlas  of  special  regions. 

SACHS,  E.  On  the  Structure  and  Functional  Relations  of  the  Optic  Thalamus.  Brain, 
Vol.  CXXVI.,  1909. 

SABIN,   F.  R.'    Atlas  of  the  Medulla  and  Midbrain.      1901. 

SCHAFER  AND  SYMINGTON.     Neurology.      In  Quain's  Anatomy,  Vol.  III.,  1908. 

SMITH,  E.  A  new  Topographical  Survey  of  the  Human  Cerebral  Cortex.  Jour,  of 
Anatomy  and  Physiology,  Vol.  XLI.,  19^07. 

STREETER,  G.  L.  Die  Entwickelung  des  Nervensystems.  In  Keibel  and  Mall's  Hand- 
buch d.  Entwickelungsgeschichte  d.  Menschen.  Bd.  II.,  1911. 

TILNEY,  F.  Contribution  to  the  Study  of  the  Hypophysis  Cerebri,  with  Especial  Refer- 
ence to  Its  Comparative  Histology.  1911. 

VOGT,  C.  La  myeloarchitecture  du  thalamus  du  cercopitheque.  Jour.  f.  Psychol.  u. 
Neurol.,  Bd.  XII.,  1909. 

WERNICKE,  C.     Atlas  des  Gehirns.      1897. 

ZIEHEN,  T.  Die  Morphologic  des  Centralnervensystems  der  Saugetiere.  In  Hertwig's 
Handbuch  der  Entwickelungslehre.  Bd.  II.,  Th.  3,  1906. 

ZIEHEN,  T.  Die  Histogenese  von  Hirn-  und  Riickenmark.  In  Hertwig's  Handbuch  der 
Entwickelungslehre.  Bd.  II.,  Th.  3,  1906. 

ZIEHEN.   T.      Nervensystem.      In  Bardeleben's  Handbuch  der  Anatomic.      1899. 


NDEX 


Abducens,  176 
Accessorius,   185 
Acervulus,  56 
Acusticus,  179 
Ala  cinerea,  82 

lobuli  centralis,  73 

uvulae,  75 
Alveus,  45,  120 
Angulus  gyri  olfact.  lat,  28 
Ansa  lenticularis,  144,  219 

peduncularis,  144,  219 
Anterior  column,  90 
tracts,  161 

commissure,  135 

ground  bundle,  161 
Apertura  medialis  ventric.  quarti  (Magendii), 

lateralis  ventric.  quarti   (Luschkae),  80 
Aphasia,  202 
Apraxia,  130 

Aquaeduct.  cerebri  Sylvii,  68 
Arachnoidea  cerebri,  88 

spinalis,  93 

Arachnoidal  villi  (Pacchioni),  88 
Arbor  medullaris,  76 

vermis  vitae,  76 
Arcuate  fibres,  167,  170 

nucleus,  84,  170 
Area  acustica,  82 

medial,  trigoni  N.  XII.,  82 

parolfactoria   (Broca),  27 

plumiformis,  82 

postrema,  82 

Ascensus  medullae  spinalis,  9 
Association  cells,  160 

area,  171 

centres   (Flechsig),  127 

conduction,   133 

fibres,  134 

tracts,  197 
Astrocytes,  106 
Astropilemma,  106 
Auditory  centre,  126 

path,  190 
Axis-cylinder,  in 

process,  109 
Axone,  109 

Baillarger's  stripe,  39,  115 
Band  of  Giacomini,  34 


Bandelette  mediale  (Gombault-Philippe),  166 
Basal  ganglia,  46 
Basket  cells,  154 
Basis  cerebri,  12 

pedunculi,  67 
Bechterew's  nucleus,  181 
Bipolar  cells,  109 
Brachia  cerebelli,  76,  156 
ad  cerebrum,  76 
ad  corp.  quadrig.,  76 
ad  medullam,  76 
ad  pontem,  76,  155 

conjunctiva,  70,   76,   156 

corp.  mamillar.,  57 

pontis,  76,  155 

quadrigemina,  66 
Brain  development,  4 

form,  9 

membranes,  86 

morphology,  3 

size,  10 

weight,  10 
Brain-sand,  56 
Brain-stem,  6 
Brain-vesicles,  4 
Broca's    callosal    stalks,   30 

centre,  128 

convolution,  20 

diagonal  band,  30 

field,  27 
Bulbus  cornu  post.,  43 

olfactorius,  27,  118 
Burdach's  column,  78,  164 

Cajal's  cells,  in,  115 
Calamus  scriptorius,  81 
Calcar  avis,  43 
Callosal  convolutions,  36 

radiations,  40 

Calloso-marginal  fissure,  23 
Capsula  externa,  48,  135 

extrema,  50,  136 

interna,  48,  61,  142 
Cauda  equina,  9 
Cavum  epidurale,  93 

interdurale,  93 

psalterii,  44 

septi  pellucidi,  41 

subdurale,  93 

281 


282 


INDEX. 


Central  tegmental  tract,  151 
Centrifugal  tracts,  131,  191 
Centripetal  tracts,  131,  186 
Centrum  medianum   (Luys),  60,  228 

semiovale   (Vieussens),  39 
Cerebellar  cortex,  154 

falx,  87 

peduncles,  inferior,  76,  170 
middle,  76,  157 
superior,  76,  156 

tent,  76 

tracts,  155 
Cerebellum,  72,  152 
Cerebral  cortex,  39 

crura,  66 

ganglia,  46 

mantle,  18 

nerves,  172 

nerves,  table,  14 

peduncles,  66 
Cerebro-spinal  fluid,  88 
Cerebrum,  n 
Chiasma  opticum,  37,  174 
Cingulum,  135,  149 
Cisterna  ambiens,  88 

cerebello'-medullaris,  88 

chiasmatis,  88 

corporis  callosi,  88 

fossae  Sylvii,  88 

interpeduncularis,  88 
Clarke's  column,  90,  159 
Claustrum,  50 
Clava,  78 

Climbing  fibres,  155 
Cochlearis,  179 
Colliculus  facialis,  82,  178 

subpinealis,  66 
Column-cells,  160 
Columna  fornicis,  44 
Columnae   griseae,   91 
Comma  bundle  of  Schultze,  166 
Commissura  anterior,  16,  135 
alba,  91 
grisea,  91 

cerebri  magna,  40 

habenularum,  55 

hippocampi,  44,  135,  146 

posterior  cerebri,  56 
medullae,  90 

supramamillaris,  225 
Commissure-cells,  160 

fibres,  135 

Confluens  sinuum,  88 
Conus  medullaris,  9,  89 

terminalis,  89 
Convolutions,  see  Gyri 
Cornu  Ammonis,  45,  120 


Cornua  ventriculi  lat.,  40,  41,  42 
Corona  radiata,  135 
Corpora  candicantia,  56,  147 

geniculata,  56 

mamillaria,  56,  147 

restiformia,  77,   170 
Corpus  album  subrotund.,  59 

callosum,  14,  40 

fornicis,  44 

geniculatum,  56 

Luys,  61 

mamillare,  56,  147 

medullare  cerebelli,  75 

parabigeminum,  271 

patellare  (Tschish),  60 

pineale,  55 

restiforme,  77,  170 

striatum,  41,  48,  144 

subthalamicum,  61,  224 

trapezoides,  179 
Cortical  cells,  39 
Cortico-bulbar  tract,  138 
Cortico-spinal  tract,  138 
Crura  cerebelli,  76,  155 
ad  cerebrum,  76 
ad  corp.  quadrig.,  76 
cerebelli  ad  medullam,  77 
ad  pontem,  76 

fornicis,  44 
Crus  fornicis,  44 
Culmen  cerebelli,  73 
Cuneus,  24 
Cuticle-plate,  3 

Declive  cerebelli,  73 

Decussation,  cerebellar  peduncles,  156 

fillet,  167 

Corel's,  274 

Meynert's,  274 

motor,  77,  138 

pyramidal,  77,  138 

sensory,  167 
Deiters'  cells,  in 

nucleus,  181 
Dendrites,  in 
Development,  brain,  4 

ependyma  cells,  103 

nerve-cells,  105 
'neuralgia,  104 

spinal  cord,  8 

spinal  ganglia,  105 
Diagonal  band  of  Broca,  30 
Diaphragma   sellae   turcicae,   88 
Diencephalon,  52,  150 

summary,  63 

Digitationes  hippocampi,  43 
Direct  cerebellar  tract,  161 
!   Direct  sensory  cerebellar  tract,  170 


INDEX. 


283 


Dura  mater  cerebri,  87 
spinalis,  93 

Edinger-Westphal  nucleus,  176 

Embolus,  83 

Eminentia  collaterale,  43 

medialis,  81 

pyramidalis,  71 

'saccularis,  38 
Encephalon,  3 
End-brain,  4,  17,  134 
Endogenous  fibres,  160 
End-plate,  37 
Ependyma  cells,   103 

nuclear  zone,   103 
Ependymium,  103 
Epicerebral  space,  88 
Eyes,  movements  of,  184 

Facialis,  178 
Facial  knee,  178 
nucleus,    178 
Falx  cerebelli,  87 
cerebri,  87 
major,  87 
minor,  87 

Fascia  dentata  (Tarini),  32 
Fasciculus  ant.  propr.,  161 
arcuatus,  135    • 

(Foville),  71 
cerebro-spinalis  ant.,  139 

lateralis,  139 
cuneatus,  77,  90,  164 
fronto-occipitalis,  135,  207 
gracilis,  77,  90,   164 
lateral,  propr.,  162 
lenticularis  (Forel),  222 
longitudinal,  dorsal.    (Schiitz),  148,  151 
inferior,  135 
medialis,  148,  153,  182 
superior,  135 
praedorsalis,  153 
mamillaris  princeps,  147 
mamillo-tegmentalis,  147 
mamillo-thalamicus,  147 
obliquus  pontis,  71 
pyramidalis,  82,  138 
•    retroflexus  (Meynert),  148 
solitarius,  185 
sulcc^marginalis,  161 
tegmento-mamillaris,  147 
thalamicus,  224 
thalarno-mamillam,    147 
uncinatus,  135 
Vicq  d'Azyr,  147 
Fasciola  cinerea,  33 
Fastigium,  79 
Fibra  pontis,  71 


Fibrae  arci  formes,  70,  155 

arcuatae,  78,  167 

ext.  dor  sales,  170,  247 
ext.  ventrales,  170,  247 
internae,  167,  242 

pontis  profundae,  82 
superficiales,  82 

propriae,  134 

terminates,  115 
Fibrillar  network,  112 
Fila  lateralia  pontis,  71 

olfactoria,  27,  118,  144 
Fillet,  lateral,  179 

mesial,  167 

Filum  terminale,  9,  89 
Fimbria,  33,  44 
Fissura  calcarina,  24 

cerebri  lat.,  18 
longitud.,  ii 
transversa,  12 

chorioidea,  42 

collateralis,  24 

hippocampi,  24 

longitudinalis  cerebri,  n 

mediana  ant.,  77,  89 

parieto-occipital.,  20,  24 

prima  (His),  26 

rhinica,  24 

Rolandi,  19 

transversa  cerebri,  12 
Flechsig's  association  centre,  127 

direct  cerebellar  tract,   161,    170 
Feltwork,  interradial,  115 

supraradial,  115 
Flocculus,  75 

accessory,  75 

peduncle  of,  75 
Folium  vermis,  74 
Foramen  caecum,  77 

diaphragmatis,  88 

interventriculare,  8,  43 

Luschkae,  80 

Magendii,  80 

Monroi,  8,  43 
Forceps,  40 

Fore-brain,  derivatives,  65 
Forel's  tegmental  decussation,  274 
Formatio  reticularis,  85 
Fornix,  44,  137,  146 

longus  (Forel),  146     . 

periphericus  (Arnold),  135,  149 

pillars  of,  44 

transversus,  44,  146 
Fossa  cerebri  lateralis  (Sylvii),  18 

interpeduncularis  (Tarini)    68     . 

mediana,  81 

rhomboidea,  81 
Fountain  decussation,  273 


284 


INDEX. 


Frenulum  veli  medullaris,  70 
Frontal  pontile  tract,  137 

lobe,  19 
Funiculus  anterior,  90,  161 

lateralis,  90,  161 

posterior,  90,  163 

separans,  82 
Fourth  ventricle,  79 

Ganglioblasts,  106 
Ganglion-strand,  105 
Ganglion  ectomamillare,  68 

habenulae,  61,  148 

interpedunculare  (Gudden),  69,  148 

profund.  mesencephali,  69 

tegmenti  dorsale,  69,  147 
Gennari's  stripe,  39 
Germ-cells,  103 
Giacomini's  band,  34 
Globus  pallidus,  48 
Glomeruli  olfactorii,  118 
Glomus  chorioideum,  44 
Glossopharyngeus,  184 
Golgi-Holmgren  canals,  113 
Golgi's  network,  112 

cells,   in 

Coil's  column,  90,  164 
Gowers'  tract,  161,  170 
Granule  layer,  cerebellum,  154 

olf.  bulb,  119 

Gratiolet's  optic  radiation,  136,  172 
Gubler's  paralysis,  143 
Gudden's  tegmental  bundle,  147 
Gustatory  centre,  126 

paths,  190 
Gyrus,  or  Gyri,  ambiens,  27 

Andreae  Retzii,  36 

angularis,  21 

centralis  ant.,  19 
post.,  21 

cerebelli,  76 

cinguli,  30 

dentatus,  32,  121,  122 

descendens  (Ecker),  21 

diagonal  is,  30 

digitati  externi,  35 

epicallosus,  34 

fasciolaris,  34,  35 

fornicatus,  25,  30,   119 

frontalis,  20 

fusiform  is,  24 

hippocampi,  32 

insulae,  22 

intralimbicus,  35 

lingualis,  24 

occipitales,  21 

olfactorio-orbitalis,  29 


Gyrus  olfact.  lateral.,  27 

medial.,  27 
orbitales,  25 
perforatus,  29 
profundi,  18 
rectus,  25 

rhinenceph.-fusiform.,  32 
rhinenceph.-lingual.,  32 
rhinenceph.-temporales,  32 
semilunaris,  27 

'subcallosus  (Zuckerkandl),  30 
subsplenialis,  34 
supramarginalis,  21 
temporales,  21 

transversi,  21 
transitivi,  18 
uncinatus,  35 

Habenula,  55 

Hearing,  cortical  centre,  126 
Helweg's  triangular  tract,  162,  171 
Hemianopsia,  174 
Hemiopia,  174 

Hemiplegia  alternans  oculomot.,  143 
facial.,  143 

completa,  142 

cruciata,  144 

incompleta,  142 
Hemisphaerium,  17 
Heschl's  convolutions,  21 
Hind-brain,  85 

Hippocampus  (cornu  Ammonis),  45,  120 
Hypoglossus,  185 
Hypophysis,  38 
Hypothalamus,  64 

Incisura  praeoccipitalis,  20 

temporalis  (Schwalbe),  24 
Indirect  sensory  cerebellar  tract,  171 
Induseum  griseum,  33,  123 

inferius,  34 
Infundibulum,  37 
Insula,  22 

Intermedius  Wrisbergi,  178 
Inter-brain,  52 
Internal  capsule,  48 
Interolivary  stratum,   117 
Interradial  feltwork,  115 
Intumescentia  cervicalis,  89 

lumbalis,  89 
Island  of  Reil,  22 
Isthmus  gyri  fornicati,  30,  31 

rhombencephali,  4,  70 

Lamina  affixa,  41 

chorioidea  ventric.   lat,  4! 
ventric.  quarti,  79 
ventric.  tertii,  57 


INDEX. 


285 


Lamina  medullaris  circumvoluta,  121 

praecommissuralis,  30 

quadrigemina,  66 

rostralis,  15 

septi  pellucidi,  41 

terminalis,  15,  37 
Laminae  medullares  cerebelli,  76 

thalami,  60 
Lancisi's  striae,  33 
Lateral  column,  go,  161 

ground  bundle,  162 

nuclei,  85,  170 

pyramidal  tract,  161 

tracts,  90,  161 

ventricle,  40 

Lattice  layer  of  thalamus,  59,  227 
Lemniscus  lateralis,  179,  190 

medialis,  167,  187 
Leptomeninx,  86 
Ligamentum  denticulatum,  94 
Limbic  lobe,  25 
Limbus  Giacomini,  34 
Limen  insulae,  28 
Lingula  cerebelli,  73 
Liquor  cerebro-spinalis,  8 
Lissauer's  marginal  zone,  163 
Lobi  cerebelli,  72,  73,  74 

insulae,  22 

Lobuli  cerebelli,  72,  73 
Lobulus  paracentralis,  24 
Lobus  frontalis,  19 

occipitalis,  21 
olfactorius,  26 
olfact.  ant.,  27 

post.,  29 
parietalis,  20 
temporalis.  21 
Locus  caeruleus,  82,  177 
Luys'  body,  60,  228 
Lyra  Davidis,  44,  135 

Mammillary  bodies,  61 
Mantle  layer,  105 
Marginal  zone,  91 
Martinotti  cells,  115 
Massa  intermedia,  55 
Medial  fillet,  149,  167,  187 
Medulla  oblongata,  77,  166  / 

spinalis,  89,  159 
Medullary  groove.   ^ 

plate,  3 

ridge,  3 

tube,  4 

Membrana  limitans,  102 
Meninges,  86 
Mesencephalon,  4,  66 

summary,  69 
Metathalamus,  63 


Metencephalon,  4,  77 

Meynert's  fountain  decussation,  153,  273 

Mid-brain,  66 

Middle  commissure,  55 

Mitral  cells,  118 

Molecular  layer,  114,  121,  122,  154 

Monakow's  nucleus,  243 

bundle,  153,  162 
Monoplegia,  145  . 
Monticulus  cerebelli,  73 
Moss  fibres,  155 
Motor  tract,  131,  138,  151,  191 

centre,  124 
Multipolar  cells,  no 
Myelencephalon,  4,  77,  166 

Nerve  process,  108 

cells,  1 06. 

Nervus  abducens,  176 
accessorius,  185 
acusticus,  179 
cochleae,    179 
facialis,  178 
glossopharyngeus,  184 
hypoglossus,  186 
intermedius  (Wrisbergi),  178 
oculomotorius,  175 
olfactorius,  172 
opticus,  172 
Sapolini,  178 
trigeminus,  176 
trochlearis,  176 
vagus,  184 
vestibuli,  181 
Wrisbergi,  178 

Neural  or  medullary  tube,  102 
Neurite,  109 
Neuroblasts,  103 
Neurofibrillse,  112 
Neuroglia,  103 
Neuroglia  cells,  103,  106 
Neurone,  109 
Nidus  avis,  75 
Nissl's  bodies,  112 
Nodulus,  75 

Nucleus  alae  cinereae,  85,  18' 
ambiguus,  85,   184 
amygdalae,  50,  145,  219 
arcuati,  84,  170 
caudatus,  47,  144 
corpor.  geniculati,  61 
mamillaris,  61,  147 
trapezoides,  83,  179 
dentatus  cerebelli,  83 
dorsalis  (Clarkii),  91,  157 
emboliformis,  83 
eminentiae  teretis,  263 
fastigii,  83 


286 


INDEX. 


Nucl.  funiculi  cuneati,  84,  167 

gracilis,  84,   167 
globosi,  84 
habenulae,  61,  148 
hypothalamicus,  61 
intercalatus  Staderini,  251 
laterales,  84,  170 
lemnisci,  70,  82,   179   . 
lenticularis,  48 
lentiformis,  48,  144 
nervorum,  see  Cerebral  Nerves,  172 
olivaris  accessor.,  84 

inferior,  84,  170 

superior,  179 
pontis,  82 

praepositus    XII,   255 
respiratorius,  196 
reticularis  lateralis,  243 

tegmenti,  83,  155 
Roller,  251 

ruber,  69,  137,  155,  227 
salivatorius,  257 
semilunaris  (Flechsig),  60,  227 
tecti,  83 
thalami,  58,  59 

Obex,  78 
Occipital  lobe,  21 
Oculomotorius,  175 
Olfactory  bulb,  27,  118 

bundle,  basal,  149 

centre,  126,  149 

nerve,  118,  172 

path,  190 

tract,  27,.  145 

tubercle,  27 
Oliva  inferior,  84,  170 

superior,  83,  179 
Operculum,  22 
Optic  radiation,  172 

tract,  172 

Optico-acoustic  reflex  tract,  153 
Opticus,  172 
Oval  bundle,  166 

Pacchionian  granulations,  88 

Pachymeninx,  86 

Pallium,  18 

Paraplegia,  143 

Parietal  lobe,  20 

Pars  mamillaris  hypothalami,  56 

optica   hypothalami,   37 
Pedunculi  cerebri,  66 
Pedunculus  corpor.  mamillaris,  147 

flocculi,  75 

Penicilli  olfactorii,  118 
Pia  mater  cerebri,  88 
spinalis,  93 


Pineal  body,  55 

Pituitary  body,  38 

Plexus  chorioideus  ventric.  lat.,  42 

ventric.  quarti,  8u 

ventric.  tertii,  58 
Pons  Varolii,  71 
Pontile  nuclei,  82 
tegmentum,  83 
tracts,  137 
Posterior  column,  163 

comma  bundle,  166 

nuclei,  167 

oval  bundle,  166 

triangular  bundle,  166 

ventral  field,  166 
Posterior  horn  cells,  159 
Post,  longitudinal  bundle,  148,  153,  182 

of  Schiitz,  148,  153,  182 
Praecuneus,  24 

Projection  fibres,  131,  135,  186 
Prosencephalon,  4 
Protoplasmic  processes,  109 
Psalterium,  44 
Pulvinar,  55 
Pupillary  reflex,  173 
Purkinje  cells,  154 
Putamen,  48 
Pyramidal  nuclei,   170 
tracts,  137,  161,  191 
Pyramids,  decussation,  77 
Pyramis  cerebelli,  74 

Quadrate  lobule,  24 

Radiatio  corpor.  callosi,  40 
striati,  144 

strio-subthalamica,  144 

strio-thalamica,  144 
Radicular  zone  of  cord,  163 
Radii,  cortical  fibres,  116 
Randschleier,    105 
Recessus  anterior,  68 

infundibuli,  38 

lateral,  ventric.  quarti,  79 

opticus,  17 

pinealis,  55 

posterior,  68 

suprapinealis,  56 

tecti,  79 

triangularis,  58 

Reflex  collaterals,  131,  166,  \#>. 
Reflex  conduction,  131,  191 
Regio  subthalamica,  6r 
Restiform  body,  158,  170 
Rhinencephalon,  25,  144 
Rhombencephalon,  4,  70 
Roof-nucleus,  83 
Rostrum,  15 
Rugae  loci  caeruleij  82 


INDEX. 


287 


Saccus  vasculosus,  38 
Sapolini's  nerve,  178 
Schultze's  comma  bundle,  166 
Schiitz's  longitudinal  bundle,  148,  153 
Sensory  centres,  126 

cerebellar  tracts,  170 
speech  centre,  199 
tracts,  186 
Septum  anterius,  93 

cervicale  intermed.,  93 
pellucidum,  16,  41 
subarachnoideale,  93 
Sinus  occipitalis,  87 
petros.  sup.,  87 
rectus,  88 
sagittal.,  87 
transversus,  87 
Smell,  cortical  centre,  126 
Speech  centres,  199 
disturbance,  202 
path,  200 
Spinal  cord,  159 
cells,  160 
development,  8 
membranes,  93 
.    tracts,  161 
Spongioblasts,  103 
Spongiopilemma,  106 
Stratum  gelatinosum,  118 

granulosum,  115,  119,  122,  154 
griseum  centrale,  68,  225 

colliculi  sup.,  69 
lacunosum,  121 
lucidum,  121 

moleculare,  114,  118,  121,  122,  154 
oriens,  121 
radiatum,  121 
reticulare,  60,  227 
zonale,  54 

Stria  alba  tuberis  (Lenhossek),  57,  146 
cornea,  41 
medullaris,  54,  148 
olfactoria  lat,  29,  145 

med.,  27,  145  ^ 
terminalis,  41 
Striae  acusticae,  81,  179 
Lancisii,  33,  40,  145 
longitudinales  (corp.  callosi),  40 
medullares  or  acusticae,  81,  179 
Stripe  of  Baillarger,  39 
sennari,  39 
Vicq  d'Azyr,  39 
Subarachnoidal  space,  86,  88 

tissue,  88 
Subdural  space,  86 
Subiculum,  45 
Subpial  space,  88 


Substantia  cortical,  cerebelli,  83,  154 

cerebri,  38,  114 
gelatinosa  centralis,  91 

Rolandi,  91 

nigra  (Sommering),  67 
perforata  ant.,  29,  145 

post.,  68 
reticularis  alba,  247 

(Arnold),  32,  121 
grisea,  241,  247 

Sulcus  or  sulci  arcuat.  rhinencephali, 
basilaris  (pontis),  71 
centralis  insulae,  22 

Rolandi,  19 
cerebelli,  72,  73,  74,  75 
chorioideus,  54 
cinguli,  23 

circularis  (Reili),  22 
corpor.  callosi,  23 
dentato-fasciolaris,  34 
digitati  externi,  35 
fimbrio-dentat.,  33 
frontales,  19 

hypothalamicus  (Monroi),  16,  55 
interdigitales,  43 
intermedius,  41 
post.,  90 

primus  (Jensen),  21 
secundus  (Eberstaller),  21 
interparietalis,  20 
lateralis  ant.,  89 

post.,  89 
limitans,  82 
median,  fornicis,  44 

fossae  rhomboid.,  81 
post.,  89 
mesencephali  lat.,  68 

med.,  68 
Monroi,  16,  55 
nervi  oculomotorii,  67 
occipitales,  21 
occipitalis  transvers.,  20,  21 
olfactorius,  25 
orbitales,  25 
paracentralis,  23 

parietal,  transvers.  (Brissaud),  21 
parolfact.  ant,  27 

post.,  26 
postcentralis,  20 
praecentralis,    19 
radiatus,  19 
semiannularis,  28 
subcallos.  med.,  30 
subparietal.,  23 
supraorbital.  (Broca),  23 
temporales,  21 
Supraradial  feltwork,  116 
Sylvian  aqueduct,  68 


288 


INDEX. 


Sylvian  fissure,  18 

fossa,  1 8 

valley,  18 
System  of  Deiters'  nucleus,  181 

Taenia  chorioidea,  42 
fimbriae,  44 
fornicis,  42 
pontis,  71 

semicircularis,  146,  214 
tecta,  33,  40 
thalami,  58 
ventriculi  quarti,  80 

Taeniae,  42 

Tapetum,  43 

Taste,  cortical  centre,  126 
paths,  189 

Tegmen  fossae  rhomboid.,  79 
•Tegmental  decussation,  153,  273 
tract,  central,  151 

Tegmentum,  67 
pontis,  83 

Tela  chorioidea  ventric.  quarti,  79 
ventric.  tertii,  52,  57 

Telencephalon,  17,  134 

internal  configuration,  38 

Teiodendrion,  109 

Temporal  lobe,  21 

Tentorium  cerebelli,  87 

Thalamencephalon,  54 

Thalamus,  54,  58 

Third  ventricle,  57 

Tigroid,  112 

Tonsilla,  75 

Torcular  Herophili,  88 

Tracts,  projection,  186 

Tract,  bulbo-thalamicus,  151,  167 
cerebello-bulbaris,  170 
cerebello-tegmentalis,  150,  156 
cerebro-spinalis,  138,  161 
cervico-lumbalis  dorsal.,  166 
corticis  ad  pontem,  137 
cortico-habenularis,  146,  148 
cortico-mamillaris,  146 
cortico-tectales,  153 
cortico-tegmentalis,   136 
cortico-thalamici,  150 
fastigio-bulbaris,  171,  259 
habenulo-peduncularis.  148 
mamillo-tegmentalis,  147 
mamillo-thalamicus,  147 
nucleo-cerebellaris,  i/o,  181 
olfacto-ammonicus,  215 
olfacto-habenularis,   148 
olfacto-mesencephalic.,  149 
olfactorius,  27,  145 
olivo-cerebellaris,  170 


Tract,  opticus,  13,  172 

peduncularis  transvers.,  68 

ponto-cerebellares,  137,  155 

rubro-reticularis,  153 

rubro-spinalis   (Monakow),  153,  162 

rubro-thalamicus,  151 

solitarius,  185 

spinalis  N.  V.,  178 

spino-cerebellaris  dorsal.   (Flechsig),  161,  170 
ventral.  (Cowers),  70,  161,  170 

spino-olivaris  (Helweg),  162,  170 

spino-tectalis,  153,  163 

spino-thalamicus,  151,  163,  169,  186 

tecto-bulbaris,  153 

tecto-cerebellares,  153 

tecto-pontinus  (Miinzer),  153,  269 

tecto-reticularis  (Pavlow),  153 

tecto-spinalis,  153,  161,  163 

tegmento-mamillaris,  147 

thalamo-corticales,  135,  150 

thalamo-habenularis,    148 

thalamo-mamillaris,  147 

thalamo-olivaris,  151,  171 

thalamo-spinalis,  151,  163 

uncinatus,  171,  259 

vestibulo-spinalis,  161,  163,  181 
Trigeminus,  176 
Trigonum  collaterale,  43 

habenulae,  55 

lemnisci,  70 

nervi  hypoglossi,  82 

olfactorium,  27 

praecommissurale,  30 

subpineale,  66 
Trochlearis,  176 
Truncus  cerebri,  8 
Tuber  cinereum,  37 

valvulae,  74 

vermis,  74 
Tuberculum  acusticum,  82,  179 

cinereum,  79 

cuneatum,  79 

mamillare  laterale,  57 

olfactorium,  27 

thalami  ant.,  59 

Uncus,  35 
Unipolar  cells,  no 
Uvula,  75 

Vagus,  184 
Vallecula  cerebelli,  72 

lateralis,  18 

Velum  interpositum,  57 
Velum  medullare  ant.,  70 
post.,  75 

terminale  (Abbey),  34,  44 


INDEX. 


289 


Vena  cerebri  interna,  58 

magna  (Galeni),  58 

chorioidea,  58 

septi  pellucidi,  58 

terminalis,  58 
Ventral  field  of  cord,  166 
Ventriculus  Arantii,  81 

lateralis,  40 

quartus,  79 

terminalis  (Krause),  91 

tertius,  57 

Verga's  ventricle,  44 
Vermis  cerebelli,  72 
Verrucae  gyri  hippocampi,  32 
Vestibularis,  181 


Vicq  d'Azyr's  stripe,  40 

bundle,  147 

Vinculum    lingulae,   73 
Visual  centre,  127 

path,   191 
Visuo-auditory  path,  196 

Weber's  paralysis,  143 
Wernekink's  commissure,  271 
Wernicke's  centre,  129 

field,  60,   229 

pupillary  reaction,  174 
Wrisberg's  nerve,  178 
Writing  centres,  202 

Zona  incerta,  224,  225 


Date  Due 

JAN  2 

H974 

fc,  . 

-^74 

SAR  i  ' 

1976 

.V  ; 

-iC'l 

if.T 

OWT^ 

OCT  2< 

)  RtO'l) 

CAT.    NO.    23   233  PRINTED    IN    U.S.A. 


D  000  207  256  9 


WL300 
V757b 
1918 
Villiger,  Emil. 

Brain  and  spinal  cord 


Villiger,  Emil. 

Brain  and  spinal  cord 


WL300 
V757b 
1918 


CALIFORNIA  COLLEGE  OF  MEDICINE  LIBRARY 

UNIVERSITY  OF  CALIFORNIA,  IRVINE 

IRVINE,  CALIFORNIA  92664 


